An update of the host range of tomato spotted wilt virus Giuseppe Parrella, Patrick Gognalons, Kahsay Gebre Selassie, C. Vovlas, Georges Marchoux

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Giuseppe Parrella, Patrick Gognalons, Kahsay Gebre Selassie, C. Vovlas, Georges Marchoux. An update of the host range of tomato spotted wilt virus. Journal of Plant Pathology, Springer, 2003, 85 (4), pp.227-264. ￿hal-02682821￿

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Distributed under a Creative Commons Attribution - ShareAlike| 4.0 International License Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Edizioni ETS Pisa, 2003 227

INVITED REVIEW AN UPDATE OF THE HOST RANGE OF TOMATO SPOTTED WILT VIRUS

G. Parrella1, P. Gognalons2, K. Gebre-Selassiè2, C. Vovlas3 and G. Marchoux2

1 Istituto per la Protezione delle Piante del CNR, Sezione di Portici, Via Università 133, 80055 Portici (NA), Italy 2 Institute National de la Recherche Agronomique, Station de Pathologie Végétale, BP 94 - 84143 Montfavet Cedex, France 3 Dipartimento di Protezione delle Piante e Microbiologia Applicata, Università degli Studi and Istituto di Virologia Vegetale del CNR, Sezione di Bari, Via G. Amendola 165/A, 70126 Bari, Italy

SUMMARY 5% nucleic acid (RNA), 70% protein, 5% carbohydrate, and 20% lipid. The genome consists of three negative or Among plant viruses, Tomato spotted wilt virus (TSWV) ambisense ssRNA species designated as S (2.9 kb), M (4.8 is considered as the most widespread and to have the kb) and L (8.9 kb). The RNAs have partially complemen- largest host-range. This virus is responsible for numerous tary terminal sequences that allow the RNAs to adopt a epidemics in different regions of the world, mainly in hor- pseudocircular or panhandle conformation (Elliott et al., ticultural and floral crops, which are often destructive and 2000). The largest RNA (L RNA) is negative sense and cause heavy economic losses. The highly polyphagous na- monocistronic; the other two RNAs (S and M RNAs) en- ture, the efficiency of virus transmission and the biological code two proteins in an ambisense arrangement. Thus, the activity of its vectors, the rapidity with which new variants viral genome codes for five proteins in total. In particular: arise, and difficulties in the control of the vectors, make (i) RNA polymerase (330-kDa) is encoded by L RNA; (ii)

TSWV one of the most feared plant viruses by growers of protein NSM, a 34-kDa protein involved in cell-to-cell agricultural crops. Preventive and integrated cultural prac- movement of the virus, is encoded by virion sense M RNA; tices such as the eradication of weed hosts able to serve as (iii) glycoproteins G1 and G2, that form the projections on virus reservoirs, combined with vector management strate- the virus particle surface and result from processing of a gies, play a crucial role in the control of the virus. Thus, 127-kDa protein, are coded for by the complementary the availability of an up-to-date list of TSWV host plants is sense M RNA; (iv) protein NSS, a 52 kDa protein with un- a potentially useful reference for researchers and farmers. known function, is encoded by the virion sense S RNA; (v) The current list of TSWV hosts consists of 1090 plants protein N, the 29 kDa structural protein that coats the ge- species belonging in 15 families of monocotyledonous nomic RNA fragment giving rise to the nucelocapsids is plants, 69 families of dicotyledonous plants and one family encoded by the complementary sense S RNA (Hull, 2002). of pteridophytes. These are enclosed in a host-derived membrane bilayer, along with an estimated 10-20 copies of the L protein, the Key words: Tomato spotted wilt virus, host range, Thrips putative RNA-dependent RNA polymerase, encoded by spp., weeds, virus control, virus disease RNA L (Hull, 2002). In nature, TSWV is transmitted by at least eight species of thrips (Mound, 1996; Ullman et al., 1997; Groves et al., INTRODUCTION 2002), in a circulative and propagative manner (Wijkamp et al., 1993; Ullman et al., 1993). Recently, factors involved First described in 1915 in Australia by Brittlebank in the determination of vector competence have been iden- (1919), Tomato spotted wilt virus (TSWV) is the type tified in Frankliniella occidentalis and Thrips tabaci (Naga- species of the genus Tospovirus in the family Bunyaviridae, ta et al., 2002). a large family of RNA viruses most of which infect verte- TSWV is readily transmitted mechanically from sap of brate and/or invertebrate hosts. TSWV is one of the most naturally infected plants. Petunia hybrida is one of the destructive plant viruses. It is estimated that worldwide it most useful diagnostic species (Allen and Matteoni, 1991) causes losses, mainly of commercial vegetable crops, of because of the rapidity with which typical brown local le- around one billion dollars annually (Scott, 2000). The eco- sion develop, usually within 2-4 days from inoculation. nomic importance of TSWV, in addition to its biological Manually inoculated Nicotiana tabacum, Nicotiana gluti- and molecular features, has made it one of the most exten- nosa and Nicotiana benthamiana develop large necrotic lo- sively studied plant viruses. cal lesions followed by systemic mosaic and necrosis, that TSWV virions are 80-120 nm diameter, spherical, en- is sometimes lethal in N. benthamiana. Cucumis sativus also veloped, and studded with surface projections composed is a reliable assay host as it develops chlorotic local lesions of two glycoproteins, G1 and G2. Virion composition is on cotyledons 4-5 days after inoculation. DAS-ELISA using polyclonal antibodies to the whole virion is commonly used for detecting TSWV isolates, Corresponding author: G. Parrella both in plant extracts (Gonzalves and Trujillo, 1986) and Fax: +39.81.7758122 E-mail: [email protected] thrips (Cho et al., 1998; Bandla et al., 1994). Differentia- 228 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 tion among virus isolates by serological methods is possible TSWV seems to be correlated with the extensive cultiva- using antisera raised to different viral antigens, such as the tion of resistant hybrids, as reported for pepper, but it has N and G structural proteins (de Avila et al., 1990; Law and also been noted that RB isolates tend to survive in weeds Moyer, 1990; Wang and Gonsalves, 1990; Adam et al., (Roggero et al., 2002). The extreme variability of TSWV 1995). TSWV-specific monoclonal antibodies detect differ- isolates, coupled with the possibility of the exchange of ge- ences among virus isolates (Sherwood et al., 1989; netic information through reassortment of genome seg- Huguenot et al., 1990; Adam et al., 1991). ments were suggested as the main causes for the apparent TSWV molecular detection has been developed using readiness of the virus to adapt so as to overcome both nat- cDNA probes (Ronco et al., 1989; Rice et al., 1990) and ri- ural and pathogen-mediated resistances (Qiu et al., 1998; boprobes (Huguenot et al., 1990), both of which have Qiu and Moyer, 1999). proved useful for the sanitary certification of plant material Different preventive strategies are effective in TSWV (Saldarelli et al., 1996). Several PCR-based methods have management. TSWV-infected hosts in vegetable-growing been developed for the specific detection of TSWV. The regions play an important role in the epidemiology of the first PCR-based assay was developed by Mumford et al. disease, thus the elimination of weed hosts is the main (1994). Immunocapture PCR and RT-PCR were developed agronomic practice to control TSWV outbreaks (Mar- by Nolasco et al. (1993) and Weekes et al. (1996), respec- choux et al., 2000). An up-to-date list of TSWV host tively. A reliable and rapid detection of TSWV from a sin- species should help in the identification of putative natural gle infected thrip by RT-PCR has recently been reported virus reservoirs and allow the eradication of potential weed by Mason et al. (2003). A very sensitive protocol for the hosts. Best (1968) provided the first list of TSWV hosts, detection and quantification of TSWV is the real-time RT- comprising 157 dicotyledonous plant species from 29 fami- PCR assay based on TaqManTM chemistry, on both “leaf lies and 6 monocotyledonous plant species from 5 families. soak” and total RNA extracts from infected plants Subsequently, many new hosts have been discovered, so (Roberts et al., 2000). that the list has been growing steadily (Cho et al., 1987; TSWV particles can be readily detected by electron mi- Berling, 1991; Sether and De Angelis, 1991; Peters and croscope observations of leaf dip preparations with or Golbach, 1998). In the last decade, extensive surveys for without gold-immunolabelling (Milne, 1993). Ultrathin new natural and experimental hosts of TSWV have identi- sectioning of infected tissues, although time-consuming, is fied a large number of new susceptible species (Gognalons a reliable technique for virus identification because of the et al., 1996, 1999). characteristic intracellular appearance and localization of In this paper we present an “update” of TSWV-suscep- virions. These accumulate in dilations of the endoplasmic tible plant species, adopting the taxonomic system pro- reticulum and mature by budding of the nucleocapsids posed by Engler and Diels (1936), modified by Bamba- through the endoplasmic reticulun membrane (Martelli cioni (1976), according to the International Code for and Russo, 1984). Botanical Nomenclature (Greuter et al., 1994). This list In general, the two main concomitant factors that make consists of 1090 plant species capable of being locally TSWV one of the most destructive and widely distributed and/or systematically infected by TSWV. The species are plant virus are the highly polyphagous nature of its vectors from a single family of the class Pteropsida (Division X - and the relative lack of host specificity of the virus. Conse- Pteridophyta, Tab. 1), 69 families of the class Dicotyledones quently, TSWV has a very large host range and its control (Division XII - Angiospermae, Tab. 2) and 15 families of remains problematic. the class Monocotyledones (Division XII - Angiospermae, Resistant varieties are useful in minimizing losses, but Tab. 3). Most of the plants listed as hosts of TSWV belong infections by field resistance-breaking (RB) TSWV isolates to the family Asteraceae (247 species), Solanaceae (172 have been reported in nominally resistant pepper hybrids species) and Fabaceae (60 species). In this list, accessions, (Roggero et al., 1999b, 2002) as well as in resistant tomato varieties, cultivars, subspecies, hybrids as well as plants hybrids (Aramburu and Marti, 2003). Nevertheless, the ge- identified only at the genus level (e.g. Aphelandra sp.) have netic status at resistance loci (i.e. heterozigosity vs ho- been considered as distinct botanical entities. The total mozigosity) could play an important role in determining number of TSWV-susceptible plant species of each family the level of resistance. The emergence of RB isolates of is reported in parenthesis near each family name.

PTERIDOPHYTA

Table 1. Class Pteropsida

Family Species Infection Reference Polypodiaceae Asplenium nidus - Adam and Kegler, 1994 (=Pteridaceae) (2) Platycerum superbum N Kaminska and Korbin, 1991

M = experimental transmission and positive in ELISA test; N = natural infections; F = transmitted by Frankliniella occidentalis; Ff = transmitted by Frankliniella fusca; T = transmitted by Thrips tabaci; - = no information given; M* = experimental transmis- sion with development of local lesions; N* = positive bioassay. Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 229

ANGIOSPERMAE Table 2. Class Dicotyledones

Family Species Infection Reference Acanthaceae (5) Acanthus mollis N Marchoux et al., 2000 Aphelandra sp. N Louro, 1996 Aphelandra squarrosa N Edwardson and Christie, 1997 Beloperone guttata N Chatzivassiliou et al., 2000 Hypoestes talniata M Marchoux et al. (unpublished)

Aizoaceae (4) Mesembryanthemum tricolor - Best, 1968 Mollugo verticillata N Groves et al., 2002 Tetragonia expansa N Rosello et al., 1994 - Edwardson and Christie, 1997

Tetragonia tetragonoides - Adam and Kegler, 1994

Amaranthaceae (16) Alternanthera sessilis - Edwardson and Christie, 1997 Amaranthus albus N Marchoux et al., 2000 Amaranthus blitum N Jorda et al., 1995 Amaranthus caudatus N Marchoux et al., 1991 F Stobbs et al., 1992

Amaranthus graecizans - Best, 1968 Amaranthus hybridus N Cho et al., 1986 Amaranthus paniculata - Sether and De Angelis, 1991 Amaranthus powellii F Stobbs et al., 1992 Amaranthus retroflexus F Stobbs et al., 1992 N Grieco et al., 2000 N Marchoux et al., 2000

Amaranthus sp. N van Os B. et al., 1993 Amaranthus spinosus N Bond et al., 1983 N Cho et al., 1989 F Stobbs et al., 2000

Amaranthus viridis N Cho et al., 1989 Celosia argentea - Adam and Kegler, 1994 Celosia cristata - Adam and Kegler, 1994 N Chatzivassiliou et al., 2000 N Marchoux et al., 2000

Celosia sp. - Edwardson and Christie, 1997 Gomphrena globosa M Allen and Matteoni, 1991 F Pocsai et al., 1998

Apiaceae Aegopodium podagraria - Adam and Kegler, 1994 (=Umbelliferae) (15) Ammi majus N Latham and Jones, 1997 Apium graveolens var. dulce N Cho et al., 1986 N Latham and Jones, 1997

Carum carvi F Stobbs et al., 1992 Chaerophyllum sp. N Chatzivassiliou et al., 1998 Chaerophyllum temulentum N Chatzivassiliou et al., 1998 Conium maculatum F Stobbs et al., 1992 Coriandrum sativum - Best, 1968 N Marchoux et al., 2000 230 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Daucus carota M Marchoux et al., (unpublished) Foeniculum vulgare M Marchoux et al., (unpublished) Hydrocotyle asiatica N Cho et al., 1987 Orlaya sp. N Chatzivassiliou et al., 1998 Petroselinum sativum f. crispum N Marchoux et al., 2000 Torilis arvensis N Lavina et al., 1996 Trachymene caerulea - Best, 1968

Apocynaceae (7) Adenium obesum - Edwardson and Christie, 1997 Adenium sp. N Verhoeven and Roenhorst, 1994 Catharanthus roseus N Tehrani et al., 1990; F Stobbs et al., 1992; N Louro, 1996; N Chatzivassiliou et al., 2000 N Marchoux et al., 2000

Nerium oleander N Louro, 1996 Pachypodium lamerei N Kaminska and Korbin, 1994 Plumeria acutiplia N Rosello et al., 1994 - Edwardson and Christie, 1997

Vinca minor N Louro, 1996

Araliaceae (5) Fatshedera lizei N* Kaminska and Korbin, 1994 Fatsia japonica N Vaira et al., 1992a N Kaminska and Korbin, 1994 N Verhoeven and Roenhorst, 1994 N Chatzivassiliou et al., 2000 N Marchoux et al., 2000

Schefflera actinophylla (=Brassaia actinophylla) N Kaminska and Korbin, 1991 N Tehrani et al., 1990

Schefflera arboricola N Tehrani et al., 1990 Trevesia palmata N Edwardson and Christie, 1997

Aristolochiaceae (3) Aristolochia clematitis N Chatzivassiliou et al., 1998 Aristolochia elegans N Rosello et al., 1994 - Edwardson and Christie, 1997

Asarum canadense F Stobbs et al., 1992

Asclepiadaceae Asclepias curassavica N Salomone et al., 2003 (=Apocynaceae) (10) Calotropis gigantica N Rosello et al., 1994 Hoya australis N Verhoeven and Roenhorst, 1998 Hoya bella N Verhoeven and Roenhorst, 1998 Hoya carnosa N Kaminska and Korbin, 1994 Hoya linearis N Verhoeven and Roenhorst, 1998 Hoya sp. N Verhoeven and Roenhorst, 1994 Oxypetalum sp. N Verhoeven and Roenhorst, 1998 Stapelia sp. N Edwardson and Christie, 1997 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 231

Family Species Infection Reference Stephanotis floribunda N Tehrani et al., 1990 N Daughtrey, 1996 N Edwardson and Christie, 1997 N Verhoeven and Roenhorst, 1998 N Chatzivassiliou et al., 2000 N Marchoux et al., 2000

Asteraceae Agyranthemum frutescens N Marchoux et al., 2000 (=Compositae) (247) Acanthospermum hispidum - Edwardson and Christie, 1997 Achillea millefolioum M Gognalons et al., 1996 Agathaea amelloides (= Felicia amelloides) N Vaira et al., 1992a

Agathaea bergeriana (= Felicia bergeriana) N Vaira et al., 1992a

Ageratum conyzoïdes M Marchoux et al. (unpublished) Ageratum houstonianum N Tehrani et al., 1990 N Router and Gitaitis, 1993 N Marchoux et al., 2000

Ageratum mexicanum N Marchoux et al., 2000 Ambrosia artemisiifolia F Stobbs et al., 1992 Ambrosia psilostachyna F Stobbs et al., 1992 Ambrosia trifida F Stobbs et al., 1992 Antennaria neglecta F Stobbs et al., 1992 Antennaria rosea M Marchoux et al. (unpublished) Anthemis arvensis M Gognalons et al., 1996 N Grieco et al., 2000

Anthemis tinctoria N Chatzivassiliou et al., 1998 Anthemis sp. N Marchoux et al., 2000 Arctium lappa N Cho et al., 1986 F Stobbs et al., 1992 F Bautista et al., 1995

Arctium minus F Stobbs et al., 1992 - Gognalons et al., 1996

Arctium tomentosum M Marchoux et al. (unpublished) Arctotheca calendula - Edwardson and Christie, 1997 N Latham and Jones, 1997 N Wilson, 1998

Argyranthemum frutescens N Verhoeven and Roenhorst, 1998 Argyranthemum sp. - Edwardson and Christie, 1997 Artemisia dracunculus - Adam and Kegler, 1994 N Marchoux et al., 2000

Artemisia vulgaris M Gognalons et al., 1996 Aster amellus N Verhoeven and Roenhorst, 1998 Aster chinensis - Edwardson and Christie, 1997 Aster cordifolius F Stobbs et al., 1992 Aster ericoides N Verhoeven and Roenhorst, 1998 Aster laterifolius F Stobbs et al., 1992 Aster novae-angliae F Stobbs et al., 1992 232 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Aster novi-belgii M Gognalons et al., 1996 Aster radula M Marchoux et al. (unpublished) Aster rotundifolius N Marchoux et al., 2000 Aster sp. - Edwardson and Christie, 1997 Aster tataricus M Marchoux et al. (unpublished) Bellis caerulescens N Marchoux et al., 2000 Bellis perennis M Gognalons et al., 1996 Bidens discoidea - Sether and De Angelis, 1991 Bidens pilosa N Cho et al., 1986 N Cho et al., 1989

Bidens radiata M Marchoux et al. (unpublished) Bidens sp. F Stobbs et al., 1992 Bidens vulgata F Stobbs et al., 1992 Brachycome iberidifoloia N Bitterlich and MacDonald, 1993 Cacalia suaveolens M Marchoux et al. (unpublished) Calendula arvensis N Lavina et al., 1996 N Grieco et al., 2000

Calendula officinalis N Tehrani et al., 1990 N Vaira et al., 1992b N Latham and Jones, 1997 N Marchoux et al., 2000

Callistephus chinensis N Tehrani et al., 1990 N Marchoux et al., 1991 F, T Lemmetty and Lindqvist, 1993 N Marchoux et al., 2000

Callistephus sp. N Daughtrey, 1996 Carduus acanthoides F Stobbs et al., 1992 Carduus crispus M Marchoux et al. (unpublished) Carduus nutans N Chatzivassiliou et al., 1998 Carduus pycnocephalus M Gognalons et al., 1996 Carthamus tinctorius N Rosello et al., 1994 Centaurea alpestris M Marchoux et al. (unpublished) Centaurea calcitrapa M Gognalons et al., 1996 Centaurea cineraria - Edwardson and Christie, 1997 Centaurea cinerea - Adam and Kegler, 1994 Centaurea collina M Marchoux et al. (unpublished) Centaurea cyanus N Sether and De Angelis, 1991 F Stobbs et al., 1992

Centaurea montana N Verhoeven and Roenhorst, 1998 Centaurea scabiosa M Marchoux et al. (unpublished) Centaurea sp. N Verhoeven and Roenhorst, 1994 Chamomilla recutita N Chatzivassiliou et al., 1998 Chamomilla suaveolens N Chatzivassiliou et al., 1998 Chondrilla juncea N Chatzivassiliou et al., 1998 M Marchoux et al. (unpublished)

Chondrilla sp. - Sether and De Angelis, 1991 Chrysanthemum carinatum M Marchoux et al. (unpublished) Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 233

Family Species Infection Reference Chrysanthemum coronarium N Cho et al., 1986 - Sether and De Angelis, 1991

Chrysanthemum frutescens N Tehrani et al., 1990 N Verhoeven and Roenhorst, 1994

Chrysanthemum indicum N Edwardson and Christie, 1997 Chrysanthemum leucanthemum F Stobbs et al., 1992 N Edwardson and Christie, 1997

Chrysanthemum macrophyllum M Marchoux et al. (unpublished) Chrysanthemum maximum - Sether and De Angelis, 1991 Chrysanthemum x morifolium N Cho et al., 1986 N Matteoni et al., 1989 N Marchoux et al., 1991 N Hausbeck et al., 1992 N Kaminska and Korbin, 1994 N Nagata et al., 1995 N Louro, 1996 N Latham and Jones, 1997

Chrysanthemum x superbum N Tehrani et al., 1990 Cichorium endivia N Verhoeven and Roenhorst, 1994 Cichorium endivia var. crispa N Marchoux et al., 2000 Cichorium endivia var. latifolia N Marchoux et al., 2000 Cichorium intybus F Stobbs et al., 1992 N Verhoeven and Roenhorst, 1994 N Louro, 1996 N Grieco et al., 2000

Cineraria cruenta (=Senecio cruentus) N Gognalons et al., 1996 Cineraria sp. N Edwardson and Christie, 1997 Cirsium acaule M Marchoux et al. (unpublished) Cirsium arvense F Stobbs et al., 1992 N Lavina et al., 1996 N Grieco et al., 2000

Cirsium japonicum N Verhoeven and Roenhorst, 1998 Cirsium lanceolatum N Edwardson and Christie, 1997 Cirsium oleraceum M Marchoux et al. (unpublished) Cirsium palustre N Van Os et al., 1993 Cirsium sp. N Verhoeven and Roenhorst, 1994 Cirsium vulgare F Stobbs et al., 1992 Conyza bonariensis N Cho et al., 1986 Conyza canadensis M, N Gognalons et al., 1996 Coreopsis drummondii N Gognalons et al., 1996 Coreopsis lanceolata F Stobbs et al., 1992 Coreopsis maritima M Marchoux et al. (unpublished) Coreopsis tinctoria - Edwardson and Christie, 1997 Cosmos bipinnatus F Stobbs et al., 1992 Cosmos sulphureus M Marchoux et al. (unpublished) Crepis alpestris M Marchoux et al. (unpublished) Crepis capillaris - Best, 1968 F Stobbs et al., 1992

Crepis divaricata - Best, 1968 234 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Crepis japonica N Edwardson and Christie, 1997 Crepis pulchra - Best, 1968 Crepis pumila - Best, 1968 Crepis rhoeadifolia - Best, 1968 Crepis rubra M Marchoux et al. (unpublished) Crepis sancta M Gognalons et al., 1996 Crepis setosa M Gognalons et al., 1996 Cynara scolymus N Vaira et al., 1992a N Vovlas and Lafortezza, 1994 N Latham and Jones, 1997 N Marchoux et al., 2000

Cynara syriaca N Gognalons et al., 1996 Dahlia arborescens M Marchoux et al. (unpublished) Dahlia hortensis N Marchoux et al., 2000 Dahlia hybrida N Marchoux et al., 1991 N Daughtrey, 1996 - Edwardson and Christie, 1997 N Latham and Jones, 1997

Dahlia pinnata x D.coccinea - Sether and De Angelis, 1991 Dahlia sp. N Hausbeck et al., 1992 N Edwardson and Christie, 1997

Dahlia variabilis N Gognalons et al., 1996 Dendranthema grandiflora N Kaminska and Korbin, 1991 Dendranthema indicum N Verhoeven and Roenhorst, 1998 Dimorphoteca sinuata N Rosello et al., 1994 N Manoussopoulos et al., 1999

Dimorphoteca sp. N Marchoux et al., 2000 Dracopis amplexicaulis N Johnson et al., 1995 Emilia javanica - Edwardson and Christie, 1997 Emilia sagittata - Edwardson and Christie, 1997 Emilia sonchifolia N Cho et al., 1986 N Gonsalves and Trujillo, 1986 N Nagata et al., 1995

Emilia sp. - Cho et al., 1987 Erigeron acer M Marchoux et al., 2003 Erigeron bonariensis N Cho et al., 1986 N Latham and Jones, 1997

Erigeron canadensis N Marchoux et al., 1991 F Stobbs et al., 1992 N Lavina et al., 1996 N Louro, 1996 N Grieco et al., 2000

Erigeron sp. N Van Os et al., 1993 Erigeron strigosus F Stobbs et al., 1992 Erlangea tomentosa N Edwardson and Christie, 1997 Eupatorium cannabinum M Gognalons et al., 1996 Eupatorium capillifolium N Groves et al., 2002 Eupatorium maculatum F Stobbs et al., 1992 Eupatorium rugosum M Marchoux et al. (unpublished) Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 235

Family Species Infection Reference Eupatorium urticifolium M Marchoux et al. (unpublished) Euryops sp. N Verhoeven and Roenhorst, 1998 Felicia fragilis M Marchoux et al. (unpublished) Felicia fructicosa N Verhoeven and Roenhorst, 1998 Felicia petiolata M Marchoux et al. (unpublished) Felicia sp. N Vaira et al., 1992c N Verhoeven and Roenhorst, 1994 N Gognalons et al., 1996

Flaveria bidentis N Edwardson and Christie, 1997 Gaillardia aristata N Cho et al., 1987 Gaillardia pulchella M Marchoux et al. (unpublished) Gaillardia sp. N Cho et al., 1987 - Sether and De Angelis, 1991

Gaillardia x grandiflora N Tehrani et al., 1990 - Sether and De Angelis, 1991

Galinsoga ciliata N Cho et al., 1986; F Stobbs et al., 1992; N Verhoeven and Roenhorst, 1998

Galinsoga parviflora N Cho et al., 1987 Galinsoga quadriradiata N Gognalons et al., 1996 Gazania sp. N Berling, 1991 N Ruter and Gitaitis, 1993 N Louro, 1996

Gazania quadriradiata N Edwardson and Christie, 1997 Gazania splendens N Marchoux et al., 2000 Gerbera hybrida - Edwardson and Christie, 1997 N Marchoux et al., 2000

Gerbera jamesonii N Kaminska and Korbin, 1991 Louro, 1996

Gerbera sp. N Verhoeven and Roenhorst, 1994 Gnaphalium purpureum N Groves et al., 2002 Gnaphalium uliginosum F Stobbs et al., 1992 Gnaphalium undulatum M Marchoux et al. (unpublished) Gynura aurantiaca N Matteoni et al., 1989 N Sether and De Angelis, 1991

Helianthus annuus N Edwardson and Christie, 1997 Helianthus giganteus M Marchoux et al. (unpublished) Helianthus orgyalis M Marchoux et al. (unpublished) Helichrysum bracteatum N Louro et al., 1996 Helichrysum sp. N Marchoux et al., 2000 Helipterum mangleesii - Cho et al., 1987 Helipterum roseum - Cho et al., 1987 Helminthia echioides N Gognalons et al., 1996 Inula sp. N Van Os et al., 1993 Inula viscosa M Gognalons et al., 1996 Lactuca canadensis F Stobbs et al., 1992 Lactuca floridana Ff Hobbs et al., 1993 236 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Lactuca perennis M Marchoux et al. (unpublished) Lactuca saligna - Adam and Kegler, 1994 Lactuca sativa N Cho et al., 1987 N Marchoux et al., 1991 N Vaira et al., 1992b N Nagata et al., 1995 N Louro, 1996 N Latham and Jones, 1997 F Roca et al., 1997

Lactuca sativa var. capitata N Cho et al., 1986 Lactuca sativa var. crispa N Cho et al., 1986 Lactuca sativa var. longifolia N Cho et al., 1986 F Bautista et al., 1995

Lactuca scariola N Cho et al., 1987 M Gognalons et al., 1996

Lactuca serriola (=Lactuca scariola) - Sether and De Angelis, 1991 F Stobbs et al., 1992 M Gognalons et al., 1996

Lactuca sp. N Edwardson and Christie, 1997 Lactuca virosa M Gognalons et al., 1996 Lagascea mollis N Rosello et al., 1994 Lapsana communis M Gognalons et al., 1996 Layia elegans N Cho et al., 1987 Ligularia kaempferi N Edwardson and Christie, 1997 Matricaria chamomilla M Gognalons et al., 1996 Matricaria maritima var. agrestis F Stobbs et al., 1992 N Marchoux et al., 2000

Matricaria matricarioides F Stobbs et al., 1992 Matricaria perforata N Marchoux et al., 2000 Melampodium sp. N Marchoux et al., 2000 Melilotus parviflora N Rosello et al., 1994 Onopordum acanthium N Gognalons et al., 1996 Onopordum illyricun N Chatzivassiliou et al., 1998 Osteospermum ecklonis N Marchoux et al., 2000 Osteospermum fruticosum N Daughtrey, 1996 Osteospermum sp. N Hausbeck et al., 1992 Picris echioides N Gognalons et al., 1996 Picris spinulosa M Gognalons et al., 1996 Pterotheca nemausensis M Gognalons et al., 1996 Pyrrhopappus carolinianus N Groves et al., 2002 Rudbeckia amplexicaulis N Edwardson and Christie, 1997 Rudbeckia hirta N Schuster and Halliwell, 1994 Rudbeckia laciniata M Marchoux et al. (unpublished) Rudbeckia nitida N Verhoeven and Roenhorst, 1998 Rudbeckia occidentalis N Marchoux et al., 2000 Rudbeckia sp. N Marchoux et al., 2000 Scorsonera laciniata M Gognalons et al., 1996 Senecio cineraria N Marchoux et al., 2000 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 237

Family Species Infection Reference Senecio cruentus N Cho et al., 1987 Senecio doria - Marchoux et al. (unpublished) Senecio erucifolius M Gognalons et al., 1996 Senecio jacobea N Cho et al., 1987 F Stobbs et al., 1992

Senecio paludosus M Marchoux et al. (unpublished) Senecio sp. N Verhoeven and Roenhorst, 1996 Senecio vulgaris N Marchoux et al., 1991 F Stobbs et al., 1992 N Latham and Jones, 1997

Senecio x hybridus N Cho et al., 1987 N Tehrani et al., 1990 - Sether and De Angelis, 1991

Sigesbeckia orientalis N Latham and Jones, 1997 Silybum marianum M Gognalons et al., 1996 Solidago sp. N Hausbeck et al., 1992 N Groves et al., 2002

Sonchus arvensis M Gognalons et al., 1996 Sonchus asper F Stobbs et al., 1992 Ff Hobbs et al., 1993 N Johnson et al., 1995 N Latham and Jones 1997

Sonchus oleraceus N Cho et al., 1986 F Stobbs et al., 1992 N Lavina et al., 1996 N Wilson, 1998 N Grieco et al., 2000

Sonchus sp. N Bond et al.,1983 Sonchus tenerrimus N Rosello et al., 1994 N Jorda et al., 1995 N Lavina et al., 1996

Spilantes oleracea N Marchoux et al., 2000 Spilantes acinella M Marchoux et al. (unpublished) Tagetes erecta - Edwardson and Christie, 1997 Tagetes minuta - Best, 1968 Tagetes patula N Tehrani et al., 1990 Tanacetum parthenium M Gognalons et al., 1996 Taraxacum officinale F Stobbs et al., 1992 M, N Gognalons et al., 1996

Taraxacum sp. N Bond et al., 1983 Tithonia rotundifolia N Ruter and Gitaitis, 1993 Tragopogon porrifolius N Dellavalle et al., 1999 Tragopogon pratensis M Gognalons et al., 1996 Troximon sp. - Best, 1968 Tussilago farfara F Stobbs et al., 1992 Verbesina alternifolia M Marchoux et al. (unpublished) Verbesina enceloides N Cho et al., 1986 N Sether and De Angelis, 1991 F Bautista et al., 1995 238 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Verbesina virginica M Marchoux et al. (unpublished) Xanthium italicum M Gognalons et al., 1996 Xanthium orientale M Marchoux et al. (unpublished) Xanthium saccharatum N Cho et al., 1987 N Sether and De Angelis, 1991

Xanthium spinosum N Chatzivassiliou et al., 1998 Xanthium strumarium F Stobbs et al., 1992 Xanthium strumosum N Edwardson and Christie, 1997 Youngia japonica - Sether and De Angelis, 1991 Zinnia elegans N Cho et al., 1987 N Tehrani et al., 1990 N Latham and Jones, 1997 N Marchoux et al., 2000

Zinnia peruviana M Marchoux et al. (unpublished)

Balsaminaceae (9) Impatiens balsamina N Thehrani et al., 1990 N Chatzivassiliou et al., 2000

Impatiens capensis F Stobbs et al., 1992 Impatiens holstii N Sether and De Angelis, 1991 Impatiens x hybridus N Hausbeck et al., 1992 Impatiens cv. New Guinea N Verhoeven and Roenhorst, 1994 N Daughtrey, 1996

Impatiens petersiana N Chatzivassiliou et al., 2000 Impatiens sp. N Louro, 1996 Impatiens sultani N Sether and De Angelis, 1991 Impatiens wallerana N Hausbeck et al., 1992 N Ruter and Gitaitis, 1993 N Tehrani et al., 1990 M Verhoeven and Roenhorst, 1994 N Daughtrey, 1996

Begoniaceae (6) Begonia elatior N Verhoeven and Roenhorst, 1998 Begonia gracilis - Edwardson and Christie, 1997 Begonia x hiemalis - Cho et al., 1987 Begonia semperflorens-cultorum N Edwardson and Christie, 1997 Begonia sp. N Hausbeck et al., 1992 N Verhoeven and Roenhorst, 1994 N Chatzivassiliou et al., 2000

Begonia x tuberhybrida - Edwardson and Christie, 1997

Berberidaceae (1) Berberis sp. - Adam and Kegler, 1994

Boraginaceae (9) Anchusa azurea N Chatzivassiliou et al., 1998 Anchusa capensis - Edwardson and Christie, 1997 Anchusa officinalis N Chatzivassiliou et al., 1998 Anchusa undulata N Chatzivassiliou et al., 1998 Cynoglossum sp. N Chatzivassiliou et al., 1998 Heliotropium europaeum N Chatzivassiliou et al., 1998 Myosotis alpestris - Best, 1968 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 239

Family Species Infection Reference Myosotis sp. F Stobbs et al., 1992 Symphytum tuberosum N Chatzivassiliou et al., 1998

Brassicaceae Alliaria officinalis F Stobbs et al., 1992 (=Cruciferae) (33) Alyssum maritimum N Lavina et al., 1996 Arabidopsis thaliana M German et al., 1995 Arabis sp. - Edwardson and Christie, 1997 Aubieta deltoides - Adam and Kegler, 1994 Barbarea vulgaris F Stobbs et al., 1992 Brassica campestris N Cho et al., 1987 Brassica napus N Van Os et al., 1993 Brassica oleracea - Edwardson and Christie, 1997 N Marchoux et al., 2000

Brassica oleracea var. botrytis - Sether and De Angelis, 1991 Brassica pekinensis - Edwardson and Christie, 1997 Brassica perviridis - Edwardson and Christie, 1997 Brassica rapa ssp. chinensis N Cho et al., 1986 Brassica rapa ssp. silvestris N Wilson, 1998 Capsella bursa-pastoris N Cho et al., 1986 F Stobbs et al., 1992 N Wilson, 1998

Cardamine hirsuta N Verhoeven and Roenhorst, 1998 Cardamine oligospermum F Stobbs et al., 1992 Cardamine sp. N Marchoux et al., 2000 Cardaria draba N Van Os et al., 1993 Cheiranthus cheiri - Edwardson and Christie, 1997 Coronopus didymus N Cho et al., 1986 Diplotaxis erucoides N Lavina et al., 1996 Diplotaxis muralis N Latham et al., 1997 Erysimum cheiranthoides F Stobbs et al., 1992 Erysimum cheiri N Marchoux et al., 2000 Hesperis matronalis F Stobbs et al., 1992 Hirschfeldia incana N Lavina et al., 1996 Lepidium draba N Lavina et al., 1996 Lunaria annua M Marchoux et al. (unpublished) Matthiola incana N Chatzivassiliou et al., 2000 Narsturtium officinale N Chatzivassiliou et al., 1998 Raphanus raphanistrum N Van Os et al., 1993 Sinapis arvensis N Chatzivassiliou et al., 1998

Buddlejaceae Buddleja davidii N Mertelik et al., 1996 (=Loganiaceae) (1)

Cactaceae (3) Opuntia sp. N Adam and Kegler, 1994 Schlumbergera bridgesii N Tehrani et al., 1990 Schlumbergera truncata N Hausbeck et al., 1991 240 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Campanulaceae (16) Campanula americana F Stobbs et al., 1992 Campanula glomerata N Edwardson and Christie, 1997 Campanula isophylla N Edwardson and Christie, 1997 Campanula latiloba N Edwardson and Christie, 1997 Campanula persicifolia N Edwardson and Christie, 1997 Campanula pyramidalis N Edwardson and Christie, 1997 Campanula rapunculoides F Stobbs et al., 1992 Campanula sp. N Verhoeven and Roenhorst, 1994 Lobelia dortmanna N Marchoux et al., 2000 Lobelia erinus N Verhoeven and Roenhorst, 1998 Lobelia ricardi N Verhoeven and Roenhorst, 1998 Lobelia sp. N Verhoeven and Roenhorst, 1994 Lobelia valida N Verhoeven and Roenhorst, 1998 Specularia perfoliata N Groves et al., 2002 Trachelium caeruleum N Marchoux et al., 2000 Trachelium sp. N Edwardson and Christie, 1997

Caprifoliaceae (5) Lonicera japonica M Mertelik et al., 1996 Sambucus nigra N, F Mertelik et al., 1996 Symphoricarpos albus F Mertelik et al., 1996 Weigela florida F Mertelik et al., 1996 Weigela florida cv. Variegata nana N Ruter and Gitaitis, 1993

Caricaceae (1) Carica papaya N Gonsalves and Trujillo, 1986

Caryophyllaceae (16) Agrostemma githago F Stobbs et al., 1992 N Daughtrey, 1996

Cerastium vulgatum F Stobbs et al., 1992 Dianthus chinensis N Chatzivassiliou et al., 2000 Dianthus sp. N Anonymous, 1991 Gypsophila elegans N Tehrani et al., 1990 N Sether and De Angelis, 1991

Gypsophila paniculata N Best, 1968 Lychnis alba F Stobbs et al., 1992 Lychnis chalcedonica - Edwardson and Christie, 1997 Lychnis coronaria N Best, 1968 Saponaria officinalis F Stobbs et al., 1992 Silene alba F Stobbs et al., 1992 N Jorda et al., 1995 N Chatzivassiliou et al., 1998

Silene sp. N Chatzivassiliou et al., 1998 Silene vulgaris N Rosello et al., 1994 Spergula arvensis F Stobbs et al., 1992 N Latham et al., 1997

Stellaria media N Cho et al., 1986 N Marchoux et al., 1991 F Stobbs et al., 1992 N Latham et al., 1997 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 241

Family Species Infection Reference Stellaria sp. N Best, 1968 N Sether and De Angelis, 1991

Chenopodiaceae (14) Beta vulgaris N Marchoux et al., 2000 Chenopodium album N Cho et al., 1986 N Cho et al., 1989 F Stobbs et al., 1992 N Louro, 1996 N Latham et al., 1997 N Grieco et al., 2000

Chenopodium amaranticolor F Stobbs et al., 1992 Chenopodium ambrosioides N Cho et al., 1986 Chenopodium capitatum - Adam and Kegler, 1994 Chenopodium foetidum - Edwardson and Christie, 1997 Chenopodium foliosum - Adam and Kegler, 1994 Chenopodium gigantospermum F Stobbs et al., 1992 Chenopodium glaucum F Stobbs et al., 1992 N Chatzivassiliou et al., 1998

Chenopodium murale N Cho et al., 1986 N Cho et al., 1989 F Stobbs et al., 1992

Chenopodium quinoa M Pocsai et al., 1998 Chenopodium urbicum - Adam and Kegler, 1994 Salsola pestifer F Stobbs et al., 1992 Spinacia oleracea N Edwardson and Christie, 1997 N Marchoux et al., 2000

Convolvulaceae (16) Calystegia sepium F Stobbs et al., 1992 N Van Os et al., 1993

Convolvulus althaeoides N Chatzivassiliou et al., 1998 Convolvulus arvensis F Stobbs et al., 1993 N Lavina et al., 1996 N Louro, 1996

Convolvulus cantabrica N Van Os et al., 1993 Convolvulus sepium F Stobbs et al., 1993 Convolvulus sp. N Best, 1968 N Van Os et al., 1993

Dichondra carolinensis M Sakimura, 1961 Ipomea batatas N Edwardson and Christie, 1997 Ipomea congesta N Cho et al., 1986 N Cho et al., 1989 F Stobbs et al., 1992

Ipomea hederacea M Sakimura, 1961 Ipomea hederifolia M Sakimura, 1961 Ipomea lacunosa F Johnson et al., 1996 Ipomea purpurea - Sether and De Angelis, 1991 N Chatzivassiliou et al., 1998

Jacquemontia tamnifolia N Schuster and Halliwell, 1994 N Johnson et al., 1995 242 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Quamoclit coccinea M Sakimura et al., 1961 Sedum sp. F Stobbs et al., 1992

Cornaceae (1) Cornus sanguinea N Chatzivassiliou et al., 1998

Crassulaceae (4) Kalanchoe blossfeldiana N Hausbeck et al., 1992 N Verhoeven and Roenhorst, 1994

Kalanchoe daigremontiana N Verhoeven and Roenhorst, 1994 Kalanchoe sp. N Tehrani et al., 1990 N Verhoeven and Roenhorst, 1994

Kalanchoe thyrsiflora N Verhoeven and Roenhorst, 1994

Cucurbitaceae (12) Citrullus lanatus - Sether and De Angelis, 1991 Citrullus vulgaris - Sether and De Angelis, 1991 Cucumis melo N Marchoux et al., 2000 Cucumis mucronata - Sether and De Angelis, 1991 Cucumis sativus M Marchoux et al., 1991 N Nagata et al., 1995

Cucurbita maxima - Sether and De Angelis, 1991 Cucurbita moschata - Sether and De Angelis, 1991 Cucurbita pepo - Sether and De Angelis, 1991 N Nagata et al., 1995

Echinocystis lobata F Stobbs et al., 1992 Lagenaria leucantha - Sether and De Angelis, 1991 Luffa cylindrica - Sether and De Angelis, 1991 Sechium edule N Edwardson and Christie, 1997

Dipsacaceae (3) Scabiosa atropurpurea - Sether and De Angelis, 1991 Scabiosa sp. N Edwardson and Christie, 1997 Dipsacus fullonum M Marchoux et al. (unpublished)

Ericaceae (1) Rhododendron sp. N Hausbeck et al., 1992

Euphorbiaceae (7) Euphorbia eritrea N Salomone et al., 2003 Euphorbia heterophylla N Johnson et al., 1995 Euphorbia peplus N Van Os et al., 1993 Euphorbia pulcherrima N Kaminska and Korbin, 1994 N Marchoux et al., 2000

Mercurialis annua N Grieco et al., 2000 Pedilanthus tithymaloides - Adam and Kegler, 1994 Ricinus communis - Sether and De Angelis, 1991

Fabaceae Arachis batizocai N Rosello et al., 1994 (=Leguminosae) (60) Arachis duranesis N Rosello et al., 1994 Arachis hypogaea N Johnson et al., 1996 Arachis monticola N Rosello et al., 1994 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 243

Family Species Infection Reference Arachis pusilla N Rosello et al., 1994 Arachis stenosperma N Rosello et al., 1994 Arachis villosulicarpa N Rosello et al., 1994 Brunfelsia sp. - Edwardson and Christie, 1997 Cajanus cajan - Edwardson and Christie, 1997 Canavalia ensiformis - Edwardson and Christie, 1997 Canavalia gladiata N Rosello et al., 1994 Cassia floribunda - Edwardson and Christie, 1997 Cassia obtusifolia N Edwardson and Christie, 1997 Cassia occidentalis F Stobbs et al., 1992 Cassia tora N Rosello et al., 1994 Cicer arietinum N EPPO/CABI, 1997 Crotalaria incana N Cho et al., 1986 Crotalaria juncea N Edwardson and Christie, 1997 Crotalaria mucronata N Cho et al., 1986 Crotalaria spectabilis N Groves et al., 2002 Cytisus scoparius N Van Os et al., 1993 Desmodium tortuosum F Johnson et al., 1996 Desmodium trifolium N Rosello et al., 1994 Desmodium uncinatum N Cho et al., 1986 Dolichos lablab - Sether and De Angelis, 1991 Dolichos uniflorus - Edwardson and Christie, 1997 Gleditschia tricanthos N Mertelik et al., 1996 Glycine max N Edwardson and Christie, 1997 Glycine soja N Golnaraghi et al., 2002 Lathyrus odoratus N Edwardson and Christie, 1997 Lathyrus sativus N Zimmer and Haber, 1992 Lens culinaris N EPPO/CABI, 1997 Lupinus albus - Edwardson and Christie, 1997 Lupinus angustifolius - Edwardson and Christie, 1997 Lupinus leucophyllus N Edwardson and Christie, 1997 Lupinus polyphyllus N Edwardson and Christie, 1997 Lupinus sp. F Stobbs et al., 1992 Lupinus subcarnosus N Shuster and Halliwell, 1994. Medicago aviculare N Wilson, 1998 Medicago lupulina F Stobbs et al., 1992 Medicago polymorpha N Cho et al., 1986 Medicago sp. N Edwardson and Christie, 1997 Melilotus officinalis N Cho et al., 1986 N Cho et al., 1989 F Stobbs et al., 1992

Phaseolus angularis - Edwardson and Christie, 1997 Phaseolus aureus N Edwardson and Christie, 1997 Phaseolus lunatus - Sether and De Angelis, 1991 Phaseolus mungo N Edwardson and Christie, 1997 Phaseolus sp. N Verhoeven and Roenhorst, 1994 244 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Phaseolus vulgaris N Cho et al., 1986 N Louro, 1996 N Marchoux et al., 2000

Pisum arvense N Edwardson and Christie, 1997 Pisum sativum F Roca et al., 1997 N Marchoux et al., 2000

Tephrosia purpurea N Edwardson and Christie, 1997 Trifolium incarnatum N Edwardson and Christie, 1997 Trifolium repens F Stobbs et al., 1992 N Lavina et al., 1996

Trifolium sp. N Bitterlich and MacDonald, 1993 N Lavina et al., 1996 N Wilson, 1998

Trifolium subterraneum - Edwardson and Christie, 1997 Vicia faba N Marchoux et al., 1991 N Louro, 1996 N Latham and Jones, 1997

Vigna mungo - Edwardson and Christie, 1997 Vigna radiata - Edwardson and Christie, 1997 Vigna unguiculata ssp. sesquipedalis - Edwardson and Christie, 1997 Vigna unguiculata ssp. unguiculata N Cho et al., 1987 (= V. sinensis) M Marcoux et al., 1991 N Verhoeven and Roenhorst, 1994 N Edwardson and Christie, 1997

Gentianaceae (6) Eustoma grandiflorum N Marchoux et al., 2000 Eustoma russellianum M Edwardson and Christie, 1997 Eustoma sp. - Sether and De Angelis, 1991 Exacum affine N Tehrani et al., 1990 Exacum officinale M Edwardson and Christie, 1997 Lisianthus sp. N Rosello et al., 1994

Geraniaceae (15) Erodium ciconium N Chatzivassiliou et al., 1998 Erodium moschatum N Wilson et al., 1998 Erodium sp. N Chatzivassiliou et al., 1998 Geranium carolinianum N Groves et al., 2002 Geranium robertianum F Stobbs et al., 1992 Geranium sp. N Sether and De Angelis, 1991 N Chatzivassiliou et al., 2000

Pelargonium crispum N She rw ood and vo n Bro e mb s e n , 1991 Pelargonium x grandiflorum N Marchoux et al., 2000 Pelargonium x hortorum N Tehrani et al., 1990 N Bitterlich and Mac Donald, 1993

Pelargonium x hortorum (2N) N Hausbeck et al., 1992 Pelargonium x hortorum (4N) N Hausbeck et al., 1992 Pelargonium limoneum N Sherw ood and vo n Broemb s e n, 1991 Pelargonium peltatum N Hausbeck et al., 1992 N Verhoeven and Roenhorst, 1994 N Louro, 1996 N Marchoux et al., 2000 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 245

Family Species Infection Reference Pelargonium sp. N Chatzivassiliou et al., 2000 Pelargonium zonale N Louro, 1996 - Edwardson and Christie, 1997

Gesneriaceae (19) Achimenes sp. N Verhoeven and Roenhorst, 1998 Aeschynanthus pulcher - Adam and Kegler, 1994 Aeschynanthus sp. N Verhoeven and Roenhorst, 1998 M Edwardson and Christie, 1997

Aeschynanthus speciosus N Verhoeven and Roenhorst, 1998 Codonanthe crassifolia N Verhoeven and Roenhorst, 1998 Columnea hirta N Verhoeven and Roenhorst, 1998 Columnea kwensis N Verhoeven and Roenhorst, 1998 Columnea sp. N Hausbeck et al., 1992 N Verhoeven and Roenhorst, 1998

Gesneria sp. - Edwardson and Christie, 1997 Gloxinia hybridum - Sether and De Angelis, 1991 Gloxinia sp. - Edwardson and Christie, 1997 Gloxinia sylvatica N Verhoeven and Roenhorst, 1998 Saintpaulia ionatha N Tehrani et al., 1990 - Sether and De Angelis, 1991 N Chatzivassiliou et al., 2000 N Marchoux et al., 2000

Saintpaulia sp. M Marchoux et al. (unpublished) Sinningia speciosa N Matteoni et al., 1989 N Bertaccini and Bellardi, 1990 F Allen and Matteoni, 1991 N Marchoux et al., 1991 N Hausbeck et al., 1992 N Verhoeven and Roenhorst, 1994 N Louro, 1996

Sinningia sp. N Marchoux et al., 1991 Sinningia tigrina - Sether and De Angelis, 1991 Sinningia speciosa N Edwardson and Christie, 1997 Streptocarpus sp. - Adam and Kegler, 1994

Hypericaceae (1) Hypericum crispum N Chatzivassiliou et al., 1998

Lamiaceae Ballota nigra N Chatzivassiliou et al., 1998 (=Labiatae) (33) Coleus blumei - Adam and Kegler, 1994 Coleus sp. - Sether and De Angelis, 1991 Lamium amplexicaule F Stobbs et al., 1992 N Jorda et al., 1995

Lamium purpurea N Chatzivassiliou et al., 1998 Lavendula vera N Marchoux et al., 2000 Leonotis nepetaefolia N Cho et al., 1986 Lycopus europaeus N Chatzivassiliou et al., 1998 Marrubium vulgare F Stobbs et al., 1992 N Latham and Jones, 1997 246 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Melissa officinalis N Chatzivassiliou et al., 1998 Mentha arvensis F Stobbs et al., 1992 N Grieco et al., 2000

Mentha microphylla N Chatzivassiliou et al., 1998 Mentha piperita - Sether and De Angelis, 1991 Mentha spicata - Sether and De Angelis, 1991 Mentha sp. N Marchoux et al., 2000 Mentha suaveolens N Chatzivassiliou et al., 1998 Moluccella laevis N Rosello et al., 1994 N Jorda et al., 1995 N Edwardson and Christie, 1997

Monarda didyma - Adam and Kegler, 1994 Monarda fistulosa F Stobbs et al., 1992 Nepeta cataria - Edwardson and Christie, 1997 Nepeta nuda N Chatzivassiliou et al., 1998 Ocimum basilicum N, M Marchoux et al., 1991 N Holcomb et al., 1999

Physostegia virginiana - Adam and Kegler, 1994 Plectranthus australis - Adam and Kegler, 1994 Prunella vulgaris N Chatzivassiliou et al., 1998 Salvia pratensis N Van Os et al., 1993 Salvia splendens N Edwardson and Christie, 1997 Salvia sp. N Marchoux et al., 2000 Salvia verbenaca N Chatzivassiliou et al., 1998 Salvia officinalis N Marchoux et al., 2000 Scutelleria sp. - Sether and De Angelis, 1991 Stachys arvensis N Cho et al., 1986 Stachys sp. - Sether and De Angelis, 1991

Malvaceae (15) Abelmoschus esculentus (=Hibiscus esculentus) M Halliwell, 1988 - Sether and De Angelis, 1991

Gossypium barbadense M Shuster and Halliwell, 1994 Gossypium hirsutum M Shuster and Halliwell, 1994 M, Ff Groves et al., 1998

Hibiscus subcarnosus N Rosello et al., 1994 Hibiscus tiliaceus M Halliwell, 1988 Hibiscus trionum F Stobbs et al., 1992 Lavatera trimestris M Marchoux et al. (unpublished) Malcomia maritima - Best, 1968 Malva neglecta F Stobbs et al., 1992 N Grieco et al., 2000

Malva nicaeensis N Cho et al., 1987 Malva parviflora N Cho et al., 1986 N Cho et al., 1989 F Stobbs et al., 1992 F Bautista et al., 1995 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 247

Family Species Infection Reference Malva rotundifolia - Edwardson and Christie, 1997 Malva sp. N Edwardson and Christie, 1997 Malva sylvestris N Lavina et al., 1996 N Wilson, 1998

Sida spinosa N Johnson et al., 1996

Melastomataceae (2) Medinilla magnifica N Kaminska and Korbin, 1994 Medinilla magnifolia N Kaminska and Korbin, 1991

Moraceae (7) Ficus benjamina N Verhoeven and Roenhorst, 1998 Ficus cyathistipula N Kaminska and Korbin, 1994 Ficus elastica cv Decora N Verhoeven and Roenhorst, 1994 Ficus elastica cv Variegata N Verhoeven and Roenhorst, 1994 Ficus lyrata N Kaminska and Korbin, 1994 Ficus spp. N Verhoeven and Roenhorst, 1994 Ficus stipulata N Lavina and Batlle, 1993

Myrsinaceae (1) Ardisia sp. N Verhoeven and Roenhorst, 1998

Nolanaceae (1) Nolana sp. N Tehrani et al., 1990

Oleaceae (3) Ligustrum vulgare N Mertelik et al., 1996 Jasminium odoratissimum N Louro, 1996 Jasminium sp. N Marchoux et al., 2000

Onagraceae (10) Clarkia amoena ssp. lindleyi - Sether and De Angelis, 1991 Clarkia sp. M Marchoux et al. (unpublished) Epilobium hirsutum N Chatzivassiliou et al., 1998 Fuschia x hybrida N Matteoni et al., 1989 - Sether and De Angelis, 1991 N Marchoux et al., 2000

Fuschia sp. N Louro, 1996 Godetia grandiflora - Edwardson and Christie, 1997 Oenothera biennis F Stobbs et al., 1992 Oenothera laciniata N Groves et al., 2002 Oenothera lamarckiana M Edwardson and Christie, 1997 Oenothera sp. N Best, 1968

Oxalidaceae (5) Oxalis acetosella N Groves et al., 2002 Oxalis corniculata N Van Os et al., 1993 N Marchoux et al., 2000

Oxalis crenata N Rosello et al., 1994 Oxalis sp. N Marchoux et al., 1991 Oxalis stricta F Stobbs et al., 1992

Paeoniaceae (1) Paeonia sp. - Holmes, 1939 - Edwardson and Christie, 1997 248 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Papaveraceae (8) Fumaria muralis N Latham and Jones, 1997 M Marchoux et al. (unpublished)

Fumaria officinalis N Chatzivassiliou et al., 1998 N Marchoux et al., 2000

Papaver dubium N Grieco et al., 2000 Papaver nudicaule N Edwardson and Christie, 1997 N Rosello et al., 1994

Papaver orientalis N Rosello et al., 1994 Papaver rhoeas N Chatzivassiliou et al., 1998 Papaver somniferum N Rosello et al., 1994 Papaver sp. F Stobbs et al., 1992

Pedaliaceae (1) Sesamum indicum N Costa and Forster, 1942 - Sether and De Angelis, 1991

Phytolaccaceae (2) Phytolacca americana - Edwardson and Christie, 1997 Phytolacca decandra - Sether and De Angelis, 1991

Piperaceae (4) Peperomia fraseri N Verhoeven and Roenhorst, 1998 Peperomia obtusifolia N Kaminska and Korbin, 1994 Peperomia rotundifolia N Verhoeven and Roenhorst, 1998 Peperomia sp. N Berling, 1991 N Verhoeven and Roenhorst, 1994

Pittosporaceae (1) Pittosporum tobira N Rosello et al., 1994

Plantaginaceae (3) Plantago lanceolata F Stobbs et al., 1992 Plantago major N Grieco et al., 2000 Plantago rugelii F Stobbs et al., 1992 N Edwardson and Christie, 1997

Plumbaginaceae (6) Limonium latifolium N Cho et al., 1986 N Louro, 1996; N Marchoux et al., 2000

Limonium perezii N Vaira et al., 1992b Limonium sinuatum N Vaira et al., 1992b N Rosello et al., 1994

Limonium sp. N Vaira et al., 1992c N Verhoever and Roenhorst, 1998

Limonium tataricum N Berling, 1991 Limonium vulgare N Latham and Jones, 1997

Polemoniaceae (1) Phlox drummondii N Edwardson and Christie, 1997

Polygonaceae (14) Fagopyrum esculentum F Stobbs et al., 1992 Fallopia convolvulus N Van Os B. et al., 1993 Persicaria maculosa N Van Os B. et al., 1993 Polygonum aviculare F Stobbs et al., 1992 N Latham and Jones, 1997 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 249

Family Species Infection Reference Polygonum convolvulus F Stobbs et al., 1992 Polygonum hydropiper - Sether and De Angelis, 1991 Polygonum lapathifolium N Verhoeven and Roenhorst, 1998 Polygonum pensylvanicum N Groves et al., 2002 Polygonum persicaria F Stobbs et al., 1992 Polygonum sp. F Stobbs et al., 1992 Rumex acetosella M Marchoux et al. (unpublished) Rumex conglomeratus N Van Os B. et al., 1993 Rumex crispus F Stobbs et al., 1992 Rumex sp. N Lavina et al., 1996

Portulacaceae (4) Claytonia perfoliata - Edwardson and Christie, 1997 Montia sp. - Best, 1968 Portulaca grandiflora N Hausbeck et al., 1992 Portulaca oleracea N Cho et al., 1986 F Stobbs et al., 1992 N Lavina et al., 1996 N Latham and Jones, 1997 N Grieco et al., 2000

Primulaceae (14) Anagallis arvensis F Stobbs et al., 1992 N Marchoux et al., 2000

Anagallis arvensis var. caerulea N Latham and Jones, 1997 Cyclamen persicum N Edwardson and Christie, 1997 N Marchoux et al., 2000

Cyclamen persicum cv. Carmen N, M, F Allen and Matteoni, 1988 Cyclamen sp. N Verhoeven and Roenhorst, 1994 Lysimachia congestiflora N Verhoeven and Roenhorst, 1998 Lysimachia consisti N Verhoeven and Roenhorst, 1998 Lysimachia nummularia N Verhoeven and Roenhorst, 1998 N Marchoux et al., 2000

Primula acaulis N Verhoeven and Roenhorst, 1998 N Marchoux et al., 2000

Primula malacoides - Best, 1968 Primula obconica - Best, 1968 N Marchoux et al., 2000

Primula sinensis - Best, 1968 Primula sp. - Best, 1968 N Hausbeck et al., 1992 N Verhoeven and Roenhorst, 1994

Primula vulgaris (=Primula acaulis) N Tehrani et al., 1990

Ranunculaceae (21) Aconitum carmichaeli N Marchoux et al., 2000 Aconitum sp. N Berling, 1991 Anemone coronaria N Lisa et al., 1990 N Tehrani et al., 1990 N Marchoux et al., 1991 - Sether and De Angelis, 1991 250 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Anemone sp. N Vaira et al., 1992 N Verhoeven and Roenhorst, 1994

Aquilegia coerulea M Edwardson and Christie, 1997 Aquilegia vulgaris F Stobbs et al., 1992 Clematis flammula N Chatzivassiliou et al., 1998 Clematis vitalba N Chatzivassiliou et al., 1998 Consolida ambigua M Marchoux et al. (unpublished) Delphinium cultorum M Marchoux et al. (unpublished) Delphinium hybridum N Latham et al., 1997 Delphinium sp. N Edwardson and Christie, 1997 N Marchoux et al., 2000

Paeonia lactiflora - Edwardson and Christie, 1997 Ranunculus abortivus F Stobbs et al., 1992 Ranunculus acris F Stobbs et al., 1992 Ranunculus arvensis F Stobbs et al., 1992 Ranunculus asiaticus N Lisa et al., 1990 N Daughtrey, 1996

Ranunculus bulbosus N Groves et al., 2002 Ranunculus hibridus N Rosello et al., 1994 Ranunculus sardous Ff Hobbs et al., 1993 N Johnson et al., 1995

Ranunculus sp. N Bond et al., 1983 N Hausbeck et al., 1992 N Vaira et al., 1992

Resedaceae (1) Reseda phyteuma M Marchoux et al. (unpublished)

Rhamnaceae (1) Rhamnus sp. N Marchoux et al., 2000

Rosaceae (7) Fragaria vesca - Sether and De Angelis, 1991 Potentilla reptans N Chatzivassiliou et al., 1998 Rubus idaeus N Edwardson and Christie, 1997 Rubus sp. N Lavina et al., 1996 Rubus ulmifolius N Chatzivassiliou et al., 1998 Sanguisorba minor N Chatzivassiliou et al., 1998 Sorbus aucuparia N Edwardson and Christie, 1997

Rubiaceae (9) Bouvardia sp. N Vaira et al., 1992 N Verhoeven and Roenhorst, 1994

Coffea arabica - Edwardson and Christie, 1997 Galium aparine N Chatzivassiliou et al., 1998 Galium lucidium N Chatzivassiliou et al., 1998 Galium parisiensis N Rosello et al., 1994 N Jorda et al., 1995

Galium sp. N Edwardson and Christie, 1997 Galium tricornutum N Rosello et al., 1994 N Jorda et al., 1995 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 251

Family Species Infection Reference Galium verum N Chatzivassiliou et al., 1998 Gardenia jasminoides N Tehrani et al., 1990 N Hausbeck et al., 1992

Saxifragaceae (5) Hydrangea macrophylla N Louro, 1996 N Marchoux et al., 2000

Hydrangea sp. N Tehrani et al., 1990 M Mertelik et al., 1996

Saxifraga sp. N Rosello et al., 1994 Saxifraga tricolor - Edwardson and Christie, 1997 Tolmiea menziesii - Adam and Kegler, 1994

Scrophulariaceae (21) Antirrhinum majus N Tehrani et al., 1990 N Vaira et al., 1992 N Latham and Jones, 1997 N Marchoux et al., 2000

Calceolaria crenatifolia - Sether and De Angelis, 1991 Calceolaria herbaceihybrida - Sether and De Angelis, 1991 Calceolaria hybrida N Marchoux et al., 2000 Calceolaria sp. N Rosello et al., 1994 Chelone sp. - Adam and Kegler, 1994 Diascia barberae N Marchoux et al., 2000 Digitalis sp. - Adam and Kegler, 1994 Linaria canadense N Groves et al., 2002 Mimulus sp. N Marchoux et al., 2000 Penstemon hartwegii N Louro, 1996 Penstemon hirsutus - Edwardson and Christie, 1997 Torenia fournieri M Marchoux et al. (unpublished) Verbascum blattaria N Chatzivassiliou et al., 1998 Verbascum thaspus F Stobbs et al., 1992 Veronica agrestis N Rosello et al., 1994 N Marchoux et al., 2000

Veronica chamaedrys N Chatzivassiliou et al., 1998 Veronica hederifolia N Rosello et al., 1994 N Marchoux et al., 2000

Veronica officinalis N Chatzivassiliou et al., 1998 Veronica persica N Chatzivassiliou et al., 1998 N Marchoux et al., 2000

Veronica sp. - Adam and Kegler, 1994

Solanaceae (172) Atropa belladonna - Edwardson and Christie, 1997 M Gognalons et al., 1999

Browallia americana - Edwardson and Christie, 1997 Browallia sp. N Tehrani et al., 1990 Browallia speciosa N Daughtrey, 1996 Browallia speciosa-major N Rosello et al., 1994 Brunfelsia calycina M Edwardson and Christie, 1997 252 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Brunfelsia sp. - Edwardson and Christie, 1997 Capsicum annuum F Allen and Matteoni, 1991 N Marchoux et al., 1991 N Vaira et al., 1992 Ff Hobbs et al., 1993 N Kaminska and Korbin, 1994 M Verhoeven and Roenhorst, 1994 N Johnson et al., 1996 N Louro, 1996 N Latham and Jones, 1997 N Roggero et al., 1999a

Capsicum baccatum M Gognalons et al., 1999 Capsicum chacoense M Gognalons et al., 1999 Capsicum chinense PI 159236 M Boiteux and Nagata, 1993 M Hobbs et al., 1994 M Gognalons et al., 1999 M Roggero et al., 1999a

Capsicum chinense PI 152225 M Hobbs et al., 1994 M Roggero et al., 1999a

Capsicum chinense var. Habanero N Momol et al., 2000 Capsicum frutescens var. Tabasco N Momol et al., 2000 Capsicum pubescens M Gognalons et al., 1999 Capsicum sp. N Lisa et al., 1990 N Vaira et al., 1992

Cestrum aurantiarum N Verhoeven and Roenhorst, 1998 Cestrum nocturum N Verhoeven and Roenhorst, 1998 Cestrum parquii - Edwardson and Christie, 1997 Cestrum purpureum N Verhoeven and Roenhorst, 1998 Cestrum rubrum N Verhoeven and Roenhorst, 1998 Cyphomandra betacea - Edwardson and Christie, 1997 Datura arborea - Sether and De Angelis, 1991 Datura ferox - Edwardson and Christie, 1997 Datura inoxia M Gognalons et al., 1999 Datura metel - Edwardson and Christie, 1997 Datura meteloides - Edwardson and Christie, 1997 Datura quercifolia M Gognalons et al., 1999 Datura sp. N Marchoux et al., 2000 Datura stramonium M Marchoux et al., 1991 F Stobbs et al., 1992 M Verhoeven and Roenhorst, 1994 F Bautista et al., 1995

Datura tatula N Berling, 1991 Datura wrightii N Berling, 1991 Dubosia leichhardtii N Edwardson and Christie, 1997 Dubosia leichhardtii x myoporoides - Edwardson and Christie, 1997 Dubosia myoporoides - Edwardson and Christie, 1997 Hyoscyamus albus M Gognalons et al., 1999 Hyoscyamus niger N Edwardson and Christie, 1997 Lycium ferocissimum - Edwardson and Christie, 1997 Lycium procissimum N Edwardson and Christie, 1997 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 253

Family Species Infection Reference Lycopersicon chilense - Gognalons et al., 1999 Lycopersicon chmielewskii - Edwardson and Christie, 1997 M Gognalons et al., 1999

Lycopersicon esculentum N Cho et al., 1986 N Matteoni et al., 1989 N Marchoux et al., 1991 N Kaminska and Korbin, 1994 N Camele et al., 1995 N Nagata et al., 1995 N Louro, 1996 N Latham and Jones, 1997 F Roca et al., 1997 N Sikora et al., 1998 N Allen et al., 2000

Lycopersicon hirsutum F Kumar et al., 1995 M Edwardson and Christie, 1997

Lycopersicon parviflorum F Kumar et al., 1995 M, F Edwardson and Christie, 1997

Lycopersicon pennellii M, F Edwardson and Christie, 1997 M Gognalons et al., 1999

Lycopersicon peruvianum M Edwardson and Christie, 1997 Lycopersicon pimpinellifolium N Edwardson and Christie, 1997 Nicandra physaloides N Cho et al., 1986 N Cho et al., 1989

Nicotiana acuminata N Edwardson and Christie, 1997 Nicotiana alata N Ruter and Gitaitis, 1993 Nicotiana angustifolia N Berling, 1991 Nicotiana arentsii M Gognalons et al., 1999 Nicotiana atropurpureum - Edwardson and Christie, 1997 Nicotiana attenuata M Gognalons et al., 1999 Nicotiana benavidesii M Gognalons et al., 1999 Nicotiana benthamiana F Allen and Matteoni, 1991 M Marchoux et al., 1991 M Verhoeven and Roenhorst, 1994

Nicotiana bigelovii - Edwardson and Christie, 1997 Nicotiana bonariensis - Best, 1968 Nicotiana calyciflora - Cho et al., 1987 Nicotiana caudigera - Edwardson and Christie, 1997 Nicotiana chinensis - Edwardson and Christie, 1997 Nicotiana clevelandii M Marchoux et al., 1991 Nicotiana cordifolia M Gognalons et al., 1999 Nicotiana cycliflora - Berling, 1991 Nicotiana debneyi - Edwardson and Christie, 1997 Nicotiana x edwardsonii - Edwardson and Christie, 1997 Nicotiana excelsior - Sether and De Angelis, 1991 Nicotiana exigua - Best, 1968 Nicotiana fragans M Gognalons et al., 1999 Nicotiana glauca - Edwardson and Christie, 1997 Nicotiana glutinosa M Marchoux et al., 1991 M Verhoeven and Roenhorst, 1994 254 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Nicotiana goodspeedii - Best, 1968 Nicotiana ingulba M Gognalons et al., 1999 Nicotiana knightiana M Gognalons et al., 1999 Nicotiana langsdorfii - Edwardson and Christie, 1997 Nicotiana linearis M Gognalons et al., 1999 Nicotiana longiflora - Edwardson and Christie, 1997 Nicotiana macrophylla - Edwardson and Christie, 1997 Nicotiana maritima - Edwardson and Christie, 1997 Nicotiana megalosiphon - Edwardson and Christie, 1997 Nicotiana miersii - Sether and De Angelis, 1991 M Gognalons et al., 1999

Nicotiana multivalvis - Sether and De Angelis, 1991 Nicotiana nesophyla M Gognalons et al., 1999 Nicotiana noctiflora M Gognalons et al., 1999 Nicotiana nudicaulis - Best, 1968 Nicotiana occidentalis M Verhoeven and Roenhorst, 1994 Nicotiana palmeri - Sether and De Angelis, 1991 M Gognalons et al., 1999

Nicotiana paniculata - Edwardson and Christie, 1997 Nicotiana petunoides M Gognalons et al., 1999 Nicotiana plumbaginifolia - Best, 1968 Nicotiana quadrivalvis - Sether and De Angelis, 1991 Nicotiana raimondii M Gognalons et al., 1999 Nicotiana repanda - Sether and De Angelis, 1991 M Gognalons et al., 1999

Nicotiana rotundifolia - Sether and De Angelis, 1991 M Gognalons et al., 1999

Nicotiana rustica M Verhoeven and Roenhorst, 1994 Nicotiana sanderae - Edwardson and Christie, 1997 Nicotiana solanifolia - Best, 1968 Nicotiana sp. N Marchoux et al., 2000 Nicotiana stocktonii M Gognalons et al., 1999 Nicotiana suaveolens - Edwardson and Christie, 1997 Nicotiana sylvestris - Edwardson and Christie, 1997 Nicotiana tabacum M Verhoeven and Roenhorst, 1994 N Marchoux et al., 2000

Nicotiana thyrsiflora M Gognalons et al., 1999 Nicotiana tomentosa - Sether and De Angelis, 1991 M Gognalons et al., 1999

Nicotiana tomentosiformis - Best, 1968 Nicotiana trigonophylla - Sether and De Angelis, 1991 M Gognalons et al., 1999

Nicotiana undulata - Best, 1968 Nicotiana veluttina - Sether and De Angelis, 1991 Nicotiana wigandioides - Best, 1968 Nolana sp. N Tehrani et al., 1990 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 255

Family Species Infection Reference Petunia hybrida N Tehrani et al., 1990 F Allen et al, 1991 M* Marchoux et al., 1991 N Hausbeck et al., 1992 M* Verhoeven and Roenhorst, 1994 N Marchoux et al., 2000

Petunia parodi M Gognalons et al., 1999 Petunia sp. N Edwardson and Christie, 1997 Petunia violacea - Sether and De Angelis, 1991 Physalis alkekengi - Edwardson and Christie, 1997 Physalis angulata - Sether and De Angelis, 1991 Physalis brasiliensis - Sether and De Angelis, 1991 Physalis floridana M* Verhoeven and Roenhorst, 1994 - Edwardson and Christie, 1997

Physalis franchetti N Rosello et al., 1994 Physalis heterophylla F Stobbs et al., 1992 Physalis hygrophylla - Sether and De Angelis, 1991 Physalis ixocarpa N Valverde et al., 1993 N Diaz-Péres and Pappu, 2000

Physalis minima N Edwardson and Christie, 1997 Physalis peruvianum (= P. peruviana) N Edwardson and Christie, 1997 Physalis pruinosa - Sether and De Angelis, 1991 Physalis pubescens - Edwardson and Christie, 1997 Salpichroa origanifolia - Edwardson and Christie, 1997 M Gognalons et al., 1999

Salpiglossis sinuata - Edwardson and Christie, 1997 Salpiglossis sp. - Sether and De Angelis, 1991 N Edwardson and Christie, 1997

Saracha saltomata M Gognalons et al., 1999 Schizanthus pinnatus - Edwardson and Christie, 1997 Schizanthus sp. N Verhoeven and Roenhorst, 1998 N Marchoux et al., 2000

Solanum aculeatisimum - Edwardson and Christie, 1997 Solanum aethiopicum M Gognalons et al., 1999 Solanum americanum F Stobbs et al., 1992 Solanum atriciplicifolium N Edwardson and Christie, 1997 Solanum atrifolium - Adam and Kegler, 1994 Solanum aviculare N Edwardson and Christie, 1997 Solanum capsicastrum N Edwardson and Christie, 1997 Solanum carolinense N Bond et al., 1983 F Stobbs et al., 1992 N Johnson et al., 1995

Solanum ciliatum - Berling 1991 Solanum dulcamara F Stobbs et al., 1992 N Grieco et al., 2000 N Marchoux et al., 2000

Solanum eleagnifolium M Gognalons et al., 1999 Solanum grandiflorum - Sether and De Angelis, 1991 256 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Solanum indicum M Gognalons et al., 1999 Solanum integrifolium M Gognalons et al., 1999 Solanum laciniatum - Berling, 1991 Solanum lycocarpum - Edwardson and Christie, 1997 Solanum mammosum N Edwardson and Christie, 1997 Solanum marginatum - Edwardson and Christie, 1997 Solanum melongena N Marchoux et al., 1991 N Louro, 1996 N Latham et al., 1997 N Parisi et al., 1998

Solanum miniatum - Edwardson and Christie, 1997 Solanum minimatum - Adam and Kegler, 1994 Solanum muricatum N Marchoux et al., 2000 Solanum nigrum N Cho et al., 1986 N Marchoux et al., 1991 F Stobbs et al., 1992 N Johnson et al., 1995 N Latham and Jones, 1997 N Wilson, 1998 N Grieco et al., 2000

Solanum nodiflorum - Edwardson and Christie, 1997 Solanum palinacanthum - Berling, 1991 Solanum pocote - Berling, 1991 Solanum pseudocapsicum - Berling, 1991 Solanum ptycanthum F Stobbs et al., 1992 Solanum quitoense N Marchoux et al., 2000 Solanum rantonnetii N Dellavalle et al., 1999 Solanum robustum - Edwardson and Christie, 1997 Solanum sanitwongsii - Edwardson and Christie, 1997 Solanum seaforthianum N Edwardson and Christie, 1997 Solanum sodomaeum - Edwardson and Christie, 1997 Solanum spinosissimum - Sether and De Angelis, 1991 Solanum triflorum - Cho et al., 1987 Solanum trifolium - Adam and Kegler, 1994 Solanum tuberosum N Nagata et al., 1995 N Louro, 1996 N Latham and Jones, 1997

Solanum viarum - Berling, 1991 Streptosolen jamesonii - Edwardson and Christie, 1997

Tropaeolaceae (2) Tropaeolum majus N Cho et al., 1986 N Cho et al., 1989 N Marchoux et al., 2000

Tropaeolum sp. - Sether and De Angelis, 1991

Ulmaceae (1) Ulmus pumila var. arborea M Mertelik et al., 1996

Urticaceae (4) Parietaria officinalis N Lavina et al., 1996 Pilea pumila F Stobbs et al., 1992 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 257

Family Species Infection Reference Urtica dioica - Edwardson and Christie, 1997 Urtica sp. N Marchoux et al., 2000

Valerianaceae (2) Centranthus ruber N Hausbeck et al., 1992 Valerianella olitoria N Vaira et al., 1992c N Marchoux et al., 2000

Verbenaceae (10) Clerodendron thomsioniae N Marchoux et al., 2000 Lantana sp. N Hausbeck et al., 1992 Verbena alternifolia M Marchoux et al. (unpublished) Verbena brasiliensis N Bond et al., 1983 Verbena hastata F Stobbs et al., 1992 Verbena hybrida - Edwardson and Christie, 1997 Verbena litoralis N Cho et al., 1986 Verbena officinalis N Chatzivassiliou et al., 1998 N Grieco et al., 2000 N Marchoux et al., 2000

Verbena rigida - Sether and De Angelis, 1991 Verbena sp. - Best, 1968 F Stobbs et al., 1992

Violaceae (5) Viola cornuta N Marchoux et al., 2000 Viola odorata M Edwardson and Christie, 1997 Viola sororia - Sether and De Angelis, 1991 Viola sp. N Marchoux et al., 2000 Viola x wittrockiana N Ruter and Gitaitis, 1993

Zygophyllaceae (1) Tribulus terrestris - Cho et al., 1987

M = experimental transmission and positive in ELISA test; N = natural infections; F = transmitted by Frankliniella occidentalis; Ff = transmitted by Frankliniella fusca; T = transmitted by Thrips tabaci; - = no information given; M* = experimental transmis- sion with development of local lesions; N* = positive bioassay.

Table 3. Class Monocotyledones

Family Species Infection Reference Agavaceae Cordyline sp. M Edwardson and Christie, 1997 (=Dracenaceae)(11) Cordyline terminalis N Cho et al., 1986 Dracaena deremensis N Kaminska and Korbin, 1994 Dracaena draco N Louro, 1996 Dracaena fragrans N Tehrani et al., 1990 Dracaena fragrans var. massangeana N Kaminska and Korbin, 1994 Dracaena marginata N Tehrani et al., 1990 Dracaena marginata bicolor N Kaminska and Korbin, 1994 Dracaena sp. N Hausbeck et al., 1992 Yucca aloifolia N Kaminska and Korbin, 1994 Yucca baccata N Tehrani et al., 1990 258 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Amaryllidaceae (30) Alstroemeria aurantiaca - Adam and Kegler, 1994 Alstroemeria hybrida N Lathman and Jones, 1997 Alstroemeria sp. N Tehrani et al., 1990 N Kaminska and Korbin, 1994 N Verhoeven and Roenhorst, 1994 N Marchoux et al., 2000

Amaryllis aulicum - Edwardson and Christie, 1997 Amaryllis belladonna N Derks and Lemmers, 1996 Amaryllis hybrida - Edwardson and Christie, 1997 Amaryllis reginae N Edwardson and Christie, 1997 Amaryllis rutilum N Edwardson and Christie, 1997 Amaryllis sp. N Edwardson and Christie, 1997 Amaryllis x nerine N Edwardson and Christie, 1997 Clivia miniata M Edwardson and Christie, 1997 Clivia sp. N Verhoeven and Roenhorst, 1998 Crinum giganteum - Edwardson and Christie, 1997 Crinum imbricatum - Edwardson and Christie, 1997 Eucharis grandiflora M Marchoux et al. (unpublished) Eucharis sp. N Verhoeven and Roenhorst, 1998 Hippeastrum aulicum - Berling, 1991 Hippeastrum x hybridum N Marchoux et al., 2000 Hippeastrum hybride N Verhoeven and Roenhorst, 1998 Hippeastrum reginae - Berling, 1991 Hippeastrum rutilum - Berling, 1991 Hippeastrum sp. N Verhoeven and Roenhorst, 1994; N Derks and Lemmers, 1996

Hymenocallis johnsonii - Edwardson and Christie, 1997 Hymenocallis occidentalis - Edwardson and Christie, 1997 Hymenocallis pauciflora - Edwardson and Christie, 1997 Hymenocallis solandriflora - Edwardson and Christie, 1997 Leptochiton quitoensis - Edwardson and Christie, 1997 Nerine bowdenii N Derks and Lemmers, 1996 Nerine sp. N OEPP, 1991 M Edwardson and Christie, 1997

Vallota sp. N Edwardson and Christie, 1997

Araceae (27) Aglaonema commutatum N Kaminska and Korbin, 1994 Aglaonema sp. M Edwardson and Christie, 1997 Anthurium andreanum - Cho et al., 1989 N Verhoeven and Roenhorst, 1994

Anthurium hookeri N Mertelik et al., 1996 Anthurium scherzerianum N Verhoeven and Roenhorst, 1994 Anthurium sp. N Verhoeven and Roenhorst, 1994 Arum maculatum N Chatzivassiliou et al., 1998 Arum palaestinum N Marchoux et al., 2000 Calla palustris - Edwardson and Christie, 1997 Calla sp. - Adam and Kegler, 1994 Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 259

Family Species Infection Reference Dieffenbachia sp. N Verhoeven and Roenhorst, 1994; N Kaminska and Korbin, 1994; N Chatzivassiliou et al., 2000

Epipremnum aureum N Kaminska and Korbin, 1991 Epipremnum pinnatum N Kaminska and Korbin, 1994 Philodendron rubra N Kaminska and Korbin, 1994 Philodendron tuxla N Kaminska and Korbin, 1994 Pothos sp. M Marchoux et al. (unpublished) Richardia africana - Sether and De Angelis, 1991 (see Zantedeschia aethiopica)

Richardia scabra - Sether and De Angelis, 1991

Spathiphyllum floribundum N Kaminska and Korbin, 1994 Spathiphyllum sp. N Verhoeven and Roenhorst, 1994 Syngonium podophyllum - Adam and Kegler, 1994 Zantedeschia aethiopica N Kaminska and Korbin, 1994 Zantedeschia albo-maculata - Berling, 1991 Zantedeschia elliottiana - Berling, 1991 Zantedeschia melanoleuca - Berling, 1991 Zantedeschia rehmannii - Berling, 1991 Zantedeschia sp. N Chatzivassiliou et al., 2000

Arecaceae Chamaedorea costaricana N Kaminska and Korbin, 1994 (=Palmae) (1)

Asparagaceae (1) Asparagus setaceus N Kaminska and Korbin, 1994

Bromeliaceae (3) Ananas comosus N Edwardson and Christie, 1997 Bromelia achmea N Verhoeven and Roenhorst, 1998 Guzmania sp. - Edwardson and Christie, 1997

Xanthorrhoeaceae Calectasia cyanea N Latham and Jones, 1997 (=Calectasiaceae) (1)

Cannaceae (1) Canna sp. - Sether and De Angelis, 1991

Commelinaceae (2) Tradescantia albiflora N Chatzivassiliou et al., 2000 Tradescantia sp. - Adam and Kegler, 1994

Cyperaceae (2) Cyperus esculentus F Stobbs et al., 1992 Cyperus rotundus N Chatzivassiliou et al., 1998

Iridaceae (7) Belamcanda chinensis N Sether and De Angelis, 1991 N, T Yamamoto and Ohata, 1997

Freesia x hybrida N Latham and Jones, 1997 N Marchoux et al., 2000

Gladiolus x grandiflorus N Latham and Jones, 1997 Gladiolus hybridum M Edwardson and Christie, 1997 260 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Family Species Infection Reference Gladiolus sp. N Derks and Lemmers, 1996 N Louro, 1996 N Marchoux et al., 2000

Iris sp. N Derks and Lemmers, 1996

Liliaceae Agapanthus sp. - Berling, 1991 (=Colchicaceae) (14) Agapanthus umbellatus M Edwardson and Christie, 1997 Allium cepa N Louro, 1996 Allium porrum M Marchoux et al. (unpublished) Allium sativum M Marchoux et al. (unpublished) Chlorophytum elatum M Marchoux et al. (unpublished) Gloriosa rothschildiana N Verhoeven and Roenhorst, 1998 Lilium longiflorum N Tehrani et al., 1990 Lilium sp. N Hausbeck et al., 1992 Lilium tigrinum - Best, 1968 N Marchoux et al., 2000

Maianthemum canadense F Stobbs et al., 1992 Ornitogalum thyrsoides N Rosello et al., 1994 Rohdea sp. - Adam and Kegler, 1994 Ruscus hypoglossum N Louro, 1996

Marantaceae (4) Maranta leuconeura var. kerchoveana N Kaminska and Korbin, 1994 Maranta sp. N Kaminska and Korbin, 1991 Maranta leuconeura var. tricolor N Kaminska and Korbin, 1994 N Verhoeven and Roenhorst, 1998

Martynia annua - Berling, 1991

Orchidaceae (4) Cymbidium sp. N Tehrani et al., 1990 Dendrobium sp. - Berling, 1991 Oncidium sp. N Hu et al., 1992 Palaenopsis sp. N Verhoeven and Roenhorst, 1998

Poaceae Avena fatua N Chatzivassiliou et al., 1998 (=Gramineae) (11) Cynodon dactylon N Jorda et al., 1995 Digitaria sanguinalis N Grieco et al., 2000 Echinocloa colonum N Rosello et al., 1994 Eleusine indica - Sether and De Angelis, 1991 Holcus lanatus N Grieco et al., 2000 Lolium perenne N Chatzivassiliou et al., 1998 Poa annua N Rosello et al., 1994 Cetaria italica M Marchoux et al. (unpublished) Cetaria verticilliata M Marchoux et al. (unpublished) Setaria viridis N Chatzivassiliou et al., 1998

M = experimental transmission and positive in ELISA test; N = natural infections; F = transmitted by Frankliniella occidentalis; Ff = transmitted by Frankliniella fusca; T = transmitted by Thrips tabaci; - = no information given; M* = experimental transmis- sion with development of local lesions; N* = positive bioassay. Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 261

REFERENCES Camele I., Rana G.L, De Stradis A., Palumbo M., 1995. Nuove epifizie del virus della bronzatura del pomodoro (TSWV) in Adam G., Kegler H., 1994. Tomato spotted wilt virus and related Basilicata e Puglia. Petria 5: 137-142. tospoviruses. Archives of Phytopathology, 28: 483-504. Chatzivassiliou E.K., Livieratos I., Jenser G., Katis N.I., 2000. Or- Adam G., Lesemann D.E., Vetten H.J., 1991. Monoclonal anti- namental plants and thrips populations associated with tomato bodies against tomato spotted wilt virus: characterization and spotted wilt virus in Greece. Phytoparasitica 28: 1-8. application. Annals of Applied Biology 118: 87-104. Chatzivassiliou E.K., Mpoumpourakas I., Drossos E., Elefthero- Adam G., Roggero P., Malavasi F., Milne R.G., Papa G., 1995. horinos I., Jenser G., Peters D., Katis N.I., 1998. A different Approach to a general tospovirus assay using antibodies to prevalence of weeds susceptible to tomato spotted wilt virus purified tomato spotted wilt tospovirus G proteins. Bulletin in tobacco and greenhouse cultivated crops in Greece. In: Pe- OEPP/EPPO Bulletin 25: 247-257. ters D. and Golbach R. (eds.). Recent progress on tospovirus research, pp. 98. Wageningen Agricultural University, Section Allen C.T., Kharboutli M.S., McAllister C.D., Coker C.M., Coop- Virology. er P.E., 2000. Thrips, weeds and tomato spotted wilt virus. Re- Cho J.J., Mau R.F.L., German T.L., Hartmann R.W., Yudin L.S., search Series of the Arkansas Agricultural Experiment Station Gonsalves D., Provvidenti R., 1989. A multidisciplinary ap- 475: 11-16. proach to management of Tomato spotted wilt virus in Allen W.R., Matteoni J.A., 1988. Cyclamen ringspot: epidemics in Hawaii. Plant Disease 73: 375-383. Ontario greenhouses caused by the tomato spotted wilt virus. Cho J.J., Mau R.F.L., Gonsalves D., Mitchell W.C., 1986. Reser- Canadian Journal of Plant Pathology 10: 41-46. voir weed hosts of tomato spotted wilt virus. Plant Disease Allen W. R., Matteoni J.A., 1991. Petunia as an indicator plant 70:1014-1017. for use by growers to monitor for thrips carryng the tomato Cho J.J., Mau R.F.L., Hamasaki R.T., Gonsalves D., 1998. Detec- spotted wilt virus in greenhouses. Plant Disease 75: 78-82. tion of tomato spotted wilt virus in individual thrips by en- Anonymous, 1991. Plantes hôtes du Tomato Spotted Wilt Virus zyme-linked immunosorbent assay. Phytopathology 78: 1348- aux Pays-Bas. Service de la Protection des Végétaux, Wa- 1352. geningen, OEPP note d'information n°520/05. Cho J.J., Mau R.F.L., Mitchell W.C., Gonzalves D., Yudin L.S., Aramburu J., Marti M., 2003. The occurrence in north-east Spain 1987. Host list of susceptible to tomato spotted wilt virus of a variant of Tomato spotted wilt virus (TSWV) that breaks (TSWV). Research Extension Series 078, 1-10. resistance in tomato (Lycopersicon esculentum) containing the Costa A.S., Forster R., 1942. Lista de hospedeiras de virus vira- Sw-5 gene. Plant Pathology 52: 407. cabeca. Bragantia 2: 83-91. Bambacioni V.M., 1976. Botanica sistematica. Liguori, Napoli, De Avila A.C., Huguenot C., Resende R.O., Kitajima E.W., Gold- Italy. bach R.W., Peters D., 1990. Serological differentiation of 20 Bandla M.D., Westcot D.M., Chenault K.D., Ulman D.E., Ger- isolates of tomato spotted wilt virus. Journal of General Virolo- man T.L., Sherwood J.L., 1994. Use of monoclonal antibody gy 71: 2801-2807. to the nonstructural protein encoded by the small RNA of Daughtrey M.L., 1996. Detection and identification of tospoviruses tomato spotted wilt tospovirus to identify viruliferous thrips. in greenhouses. Acta Horticulturae 431: 90-105. Phytopathology 84: 1247-1431. Diaz-Pérez J.C., Pappu H.R., 2000. First report of tomato spot- Bautista R.C., Mau R.F.L., Cho R.J., Custer D., 1995. Potential of ted wilt virus infection of tomatillo in Georgia. Plant Disease tomato spotted wilt tospovirus plant hosts in Hawaii as virus 84: 1155. reservoirs for transmission by Frankliniella occidentalis Dellavalle G., Ciuffo M., Roggero P., Lisa V., Minuto A., Minuto (Thysanoptera: Thripidae). Phytopathology 85: 953-958. G., Rapetti S., 1999. Infezioni da Tospovirus in Delfinio, Berling A., 1991. Le TSWV, un fléau qui se répand. Symptômes Gazania, Margherita, Sedano, Scorzobianca e Solanum ranton- et plantes hôtes. Phytoma 431: 43-45. netii in Liguria. Informatore Fitopatologico 49: 63-64. Bertaccini A., Bellardi M.G., 1990. Tomato spotted wilt virus in- Derks A.F.L.M., Lemmers M.E.C., 1996. Detection of tospovirus- fecting gloxinia in Italy. Phytopathologia Mediterranea 29: 205- es in bulbose crops and their transmissibility by vegetative 208. propagation. Acta Horticulturae 432: 132-139. Best R.J., 1968. Tomato spotted wilt virus. Advanced Virus Re- Edwardson J.R., Christie R.G., 1997. Tospovirus. In: Virus infect- search 13: 65-146. ing pepper and other solanaceous crops. Vol. 2, pp. 687-719, University of Florida, Agricultural Experiment Station. Boiteux L.S., Nagata T., 1993. Susceptibility of Capsicum chi- Gainesville, USA. nense PI159236 to tomato spotted wilt virus isolates in Brazil. Plant Disease 77: 210. Elliott R.M., Bouloy C., Calsher C.H., Goldbach R., Moyer J.T., Nichol S.T., Petterson R., Plyusnin A., Schmaljohn C.S., 2000. Bond W.P., Whitman H.K., Blanck L.L., 1983. Indigenous weeds Family Bunyaviridae. In: Van Regenmortel M.H.V., Fauquet as reservoirs of tomato spotted wilt virus in Louisiana. Phy- C.M., Bishop D.H.L, Carstens E.B., Estes M.K., Lemon S.M., topathology 73: 499. Maniloff J., Mayo M.A., McGeoch D.J., Pringle C.R, Wickner Brittlebank C.C., 1919. Tomato diseases. Journal of Department of R.B. (eds.). Virus Taxonomy. VIIth Report of the International Agriculture of Victoria Australia 17: 231-235. Committee on Taxonomy of Viruses, pp. 599-621. Academic Press, San Diego, USA. Bitterlich I., MacDonald L.S., 1993. The prevalance of tomato spot- ted wilt virus in weeds and crops in southwestern British Colum- Engler A., Diels L., 1936. Syllabus der Planzenfamilien. Gebrun- bia. Canadian Plant Disease Survey 73: 137-142. der Borutraeger, Berlín, Germany. 262 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

EPPO/CABI, 1997. Tomato spotted wilt tospovirus. In: Quaran- Hu J.S., Wang M., Ferreira S., Ogata D., 1992. Tomato spotted tine pests for Europe, 2nd ed., pp. 1379-1387. CAB Interna- wilt virus on Oncidium orchids in Hawaii. Plant Disease 76: tional, Wallingford, U.K. 426. German, T.L., Adkins S., Witherell A., Richmond K.E, Knaack Huguenot C., Dobbelsteen G., Van Den Haan P., De Wagemak- W.R., Willis D.K., 1995. Infection of Arabidopsis thaliana eco- ers C.A.M., Drost G.A., Osterhaus A.D.M.E., Peters D., type Columbia by tomato spotted wilt virus. Plant Molecular 1990. Detection of tomato spotted wilt virus using monoclon- Biology Reporter 13: 110-117. al antibodies and riboprobes. Archives of Virology 100: 47-62. Gibbs A.J., 1990. Tomato spotted wilt tospovirus. Plant Viruses Hull R., 2002. Matthews' Plant Virology. Fourth edition, Acade- Online (http://image.fs.uidaho.edu/vide/descr837.htm) mic Press , UK. Johnson R.R., Black L.L., Hobbs H.A., Valverde R.A., Story Gognalons P., Gebre-Selassie K., Marchoux G., 1996. Le tomato R.N., Bond W.P., 1995. Association of Frankliniella fusca and spotted wilt virus dans le sud de la France. Phytoma 489: 52-56. three winter weeds with tomato spotted wilt virus in Gognalons P., Gebre-Selassie K., Marchoux G., 1999. La gamme Louisiana. Plant Disease 79: 572-576. d'hôtes du TSWV continue a s'étendre. De nuvelles Johnson, W.C., Todd J.W., Culbreath A.K., Mullinix B.G., 1996. Solanacées sont touchées. Phytoma 512: 47-50. Role of warm-season weeds in spotted wilt epidemiology in Golnaraghi A.R., Shahraeen N., Pourrahim R., Farzadfar S., the southeastern coastal plain. Agronomy Journal 88: 928-933. Ghasemi A., 2003. First report of the natural occurrence of Jorda C., Ortega A., Juarez M., 1995. New hosts of tomato spot- eight viruses affecting soybeans in Iran. New Disease Report, ted wilt virus. Plant Disease 79: 538. (http://www.bspp.org.uk/ndr/july2002/2002-7.asp) Kaminska M., Korbin M., 1991. The occurrence of tomato spot- Gonsalves D., Trujillo E., 1986. Tomato spotted wilt virus in pa- ted wilt virus in Polish greenhouses. Phytopathologica Polonica paya and detection of the virus by ELISA. Plant Disease 70: 12: 9-14. 501-506. Kaminska M., Korbin M., 1994. New natural hosts of tomato Greuter W., Barrie F., Burdet H.M., Chaloner W.G., Demoulin spotted wilt virus. Acta Horticulturae, 377: 123-128. V., Hawksworth D.L., Jorgensen P.M., Nicolson D.H., Silva Kumar N.K.K., Ullman D.E., Cho J.J., 1995. Frankliniella occi- P.C., Trehane P., McNeill J., 1994. International Code of dentalis (Thysanoptera: Thripidae) landing and resistance to Botanical Nomenclature (Tokyo Code). Regnum Vegetabile tomato spotted wilt tospovirus among Lycopersicon accessions No. 131. Koeltz Scientific Books, Koenigstein. with additional comments on Thrips tabaci (Thysanoptera: Grieco P.D., Conte D., Munno I., Nuzzaci M., De Stradis A., 2000. Thripidae). Environmental Entomology 24: 513-520. Presenza del virus della bronzatura del pomodoro (TSWV) su Latham L.J., Jones R.A. C., 1997. Occurrence of tomato spotted piante spontanee e infestanti nell'area del metapontino in Basili- wilt tospovirus in native flora, weeds, and horticultural crops. cata. Informatore Fitopatologico 50: 43-46. Australasian Journal of Agriculture Research 48: 359-369. Groves R.L., Kennedy G.G., Walgenbach J.F., Moyer J.W., 1998. Lavina A., Aramburu A., Moriones E., 1996. Occurrence of Inoculation of Tomato spotted wilt virus into cotton. Plant tomato spotted wilt and cucumber mosaic viruses in field- Disease 82: 959. grown tomato crops and associated weeds in northeastern Groves R.L., Walgenbach J.F., Moyer J.W., Kennedy G.G., 2002. Spain. Plant Pathology 45: 837-842. The role of weed hosts and tobacco thrips, Frankliniella fusca, Lavina A., Batlle A., 1993. First report of tomato spotted wilt in the epidemiology of tomato spotted wilt virus. Plant Disease virus infection of Ficus species in Spain. Plant Disease 77: 536. 86: 573-582. Law M.D., Moyer J.W., 1990. A tomato spotted wilt-like virus Halliwell R.S., 1988. Tomato spotted wilt virus infection of culti- with a serologically distinct N protein. Journal of General Vi- vated malvaceous sp. and other crop plants in south Texas. rology 71: 933-938. Phytopathology 78: 627. Lemmetty A., Lindqvist I., 1993. Thrips tabaci Lind. (Thysanoptera Hausbeck M.K., Gildow F.E., 1991. Report of tomato spotted thripidae), another vector for tomato spotted wilt virus in Fin- wilt virus on thanksgiving cactus. Plant Disease 75: 215. land. Agricultural Science in Finland 2: 189-194. Lisa V., Vaira A.M., Milne R.G., Luisoni E., Rapetti S., 1990. Hausbeck M.K., Welliver R.A., Derr M.A., and Gildow F.E. Tomato spotted wilt in cinque specie coltivate in Liguria. In- 1992. Tomato spotted wilt virus survey among greenhouse or- formatore Fitopatologico 40: 34-41. namentals in Pennsylvania. Plant Disease 76: 795-800. Louro D., 1996. Detection and identification of tomato spotted Hobbs H.A., Black L.L., Story R.N., Valverde R.A., Bond W.P., wilt virus and impatiens necrotic spot virus in Portugal. Acta Gatti J.M., Scaffer D.O., Johnson R.R., 1993. Transmission of Horticulturae 431: 99-105. tomato spotted wilt virus from pepper and three weed hosts by Frankliniella fusca. Plant Disease 77: 797-799. Manoussopoulos I.N., Chatzivassiliou E.K., Smyrnioudis I.N., Katis N.I., 1999. Two diseases of Dimorphotheca caused by Hobbs H.A., Black L.L., Johnson R.R, Valverde R.A., 1994. Dif- lettuce mosaic potyvirus and tomato spotted wilt tospovirus. ferences in reactions among tomato spotted wilt virus isolates Phytoparasitica 27: 227-232. to three resistant Capsicum chinense lines. Plant Disease 78: Mason G., Roggero P., Tavella L., 2003. Detection of Tomato 1220. spotted wilt virus in its vector Frankliniella occidentalis by re- Holcomb, G.E., Valverde R.A., Sim J., Nuss J., 1999. First report verse trascription-polymerase chain reaction. Journal of Viro- on natural occurrence of tomato spotted wilt tospovirus in logical Methods 109: 69-73. basil (Ocimum basilicum). Plant Disease 83: 966. Marchoux G., Gebre-Selassie K., Villevieille M., 1991. Detection Holmes F.O., 1939. Handbook of Phytopathogenic Viruses. of tomato spotted wilt virus and transmission by Frankliniella Burgess, Minneapolis, USA. occidentalis in France. Plant Pathology 40: 347-351. Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264 Parrella et al. 263

Marchoux G., Hostachy B., Gebre-Selassie K., Gognalons P., isolates of tomato spotted wilt tospovirus from Spain. Plant 2000. Tomato spotted wilt virus: hôtes et méthodes de lutte. Pathology 46: 407-415. PHM - Revue Horticole 418: 46-52. Roggero P., Dellavalle G., Ciuffo M., Pennazio S., 1999. Effects Martelli G.P., Russo M., 1984. Use of thin sectioning for visuali- of temperature on infection in Capsicum sp. and Nicotiana zation and identification of plant viruses. Methods in Virology benthamiana by impatiens necrotic spot tospovirus. European 8: 143-224. Journal Plant Pathology, 105: 509-512. Mertelík J., Götzová B., Mokrá V., 1996. Epidemiological aspects Roggero P., Melani V., Ciuffo M., 1999. Two field isolates of of tomato spotted wilt virus infection in the Czech republic. Tomato spotted wilt virus overcome hypersensitive response Acta Horticulturae 432: 368-375. of a pepper (Capsicum annuum) Hybrid with resistance intro- gressed from C. chinense PI152225. Plant Disease 83: 965. Milne R.J., 1993. Electron microspcopy of in vitro preparations. In: Matthews R.E.F. (ed.) Diagnosis of Plant Virus Diseases, Roggero P., Masenga V., Tavella L., 2002. Field isolates of Tomato pp. 215-251. CRC Press, Boca Raton, USA. spotted wilt virus overcoming resistance in pepper and their spread to other hosts in Italy. Plant Disease 86: 950-954. Momol M.T., Pappu H.R., Dankers W., Rich J.R., Olson S.M., Rosello S., Jordá C., Nuez F., 1994. El virus del bronceado de to- 2000. First report of tomato spotted wilt virus in habanero mate (TSWV). I. Enfermeda des y epidemiologia. Phytoma Es- and tabasco peppers in Florida. Plant Disease 84: 1154. paña 62: 21-33. Mound L.A., 1996. The Thysanoptera vector species of Ronco A.E., Dal Bo E., Ghiringhelli P.D., Medrano C., Ro- Tospovirus. Acta Horticulturae 431: 298-309. manowski V., Sarachu A.N., Grau O., 1989. Cloned cDNA Mumford R.A., Barker I., Wood K.R., 1994. The detection of probes for the detection of tomato spotted wilt virus. Phy- tomato spotted wilt virus using the polymerase chain reaction. topathology 79: 1309-1313. Journal of Virological Methods 46: 303-311. Ruter J.M., Gitaitis R.D., 1993. First report of tomato spotted Nagata T., de Ávila A.C., Barbosa C.J., Melo P.C.T., Juliatti F.C., wilt virus on bedding plants in Georgia. Plant Disease 77: 101. Kitajima E.W., 1995. Occurrence of different tospoviruses in Saldarelli P., Barbarossa L., Grieco F., Gallitelli D., 1996. Digoxi- six states of Brazil. Fitopatologia Brasiliera 20: 90-95. genin-labelled riboprobes applied to phytosanitary certifica- tion of tomato in Italy. Plant Disease 80: 1343-1346. Nagata T., Inoue-Nagata A.K., van Lent J., Goldbach R., Peters D., 2002. Factors determining vector competence and speci- Salomone A., Masenga V., Minuto G., Parodi C., Roggero P., ficity for transmission of Tomato spotted wilt virus. Journal of 2003. First report of Tomato spotted wilt virus (Tospovirus, General Virology 83: 663-671. Bunyaviridae) infecting Euphorbia eritrea and Asclepias curas- savica in Liguria, Italy. New Disease Report, (http://www.bspp. Nolasco G., de Blas C., Torres V., Ponz F., 1993. A method com- org.uk/ndr/july2003/2003-33.asp). bining immunocapture and PCR amplification in a microtiter plate for the detection of a plant viruses and subviral Schuster G.L., Halliwell R.S., 1994. Six new hosts of tomato pathogens. Journal of Virological Methods 45: 201-218. spotted wilt virus in Texas. Plant Disease 78: 100. Scott A., 2000. Tomato spotted wilt virus – positive steps towards OEPP, 1991. Plantes hôtes du tomato spotted wilt virus aux negative success. Molecular Plant Pathology 1: 151-157. Pays-Bas. Service de la Protection des Végétaux, Wageningen, OEPP note d'information n° 520/05. Sether D.M., De Angelis J.D., 1991. Tomato spotted wilt virus host list and bibliography. Agricultural Experimental Station, Parisi B., Roggero P., Grassi G., Masenga V., 1998. Tospovirus Special Report n° 888, pp. 16. Department of Entomology, (TSWV) su melanzana in coltura protetta. Informatore Agrario Oregon State University, Corvallis. 48: 45-48. Sherwood J.L., Sanborn M.R., Keyser G.C., Myers L.D., 1989. Peters D., Goldbach R., 1998. An updated list of plant species Use of monoclonal antibodies in detection of tomato spotted susceptible to tospovirus. Wageningen Agricultural University, wilt virs. Phytopathology 79: 61-64. Section Virology, The Netherlands. Sherwood J.L., von Broembsen S.L., 1991. First report of Tomato Pocsai E., Horváth J., Kazinczi G., 1998. New results on the oc- spotted wilt virus in greenhouse crops in Oklahoma. Plant currence of plant viruses in Hungarian rivers and lakes. Acta Disease 75: 750. Botanica Croatica 57: 1-10. Sikora E.J., Gudauskas R.T., Murphy J.F., Porch D.W., Andriani- Qiu W.P., Geske S.M., Hickey C.M., Moyer J.W., 1998. Tomato fahanana M., Zehnder G.W., Bauske E.M., Kemble J.M., spotted wilt tospovirus genome reassortment and genome seg- Lester D.F., 1998. A multivirus epidemic of tomatoes in Al- ment-specific adaptation. Virology 244: 186-194. abama. Plant Disease 82:117-120. Stobbs L.W., Broadbent A.B., Allen W.R., Stirling A.L., 1992. Qiu W.P., Moyer J.W., 1999. Tomato spotted wilt tospovirus Transmission of tomato spotted wilt virus by the western adapts to the TSWV N gene-derived resistance by genome re- flower thrips to weeds and native plants found in southern assortment. Phytopathology 89: 575-582. Ontario. Plant Disease 76: 23-29. Rice D., German T.L., Mau R.F.L., Fujimoto F.M., 1990. Dot blot Tehrani B., Allen W.R., Matteoni J.A., 1990. Update on the inci- detection of tomato spotted wilt virus RNA in plant and dence of tomato spotted wilt virus in greenhouse. Canadian thrips tissues by cDNA clones. Plant Disease 74: 274-276. Plant Disease Survey 70: 102-103. Roberts C.A., Dietzgen R.G., Heelan L.A., Maclean D., 2000. Re- Ullman D.E., German T.L., Sherwood J.L., Westcot D.M., Can- al-time RT-PCR fluorescent detection of tomato spotted wilt tone F.A., 1993. Tospovirus replication in insect vector cells: virus. Journal of Virological Methods 88: 1-8. immunocytochemical evidence that the nonstructural protein Roca E., Aramburu J., Moriones E., 1997. Comparative host re- encoded by the S RNA of tomato spotted wilt tospovirus is actions and Frankliniella occidentalis transmission of different present in thrips vectors cells. Phytopathology 83: 456-463. 264 Host range of TSWV Journal of Plant Pathology (2003), 85 (4, Special issue), 227-264

Ullman D.E., Sherwood J.L., German T.L., 1997. Thrips as vectors Progress in Tospovirus Research, pp. 77-80. Wageningen of plant pathogens. In: Lewis, T.L.(ed.). Thrips as crop pests, Agricultural University, Section Virology, Wageningen, The pp. 539-565. CAB International, Wallingford, U.K., Netherlands. Vaira A.M., Gallo S., Lisa V., 1992a. Nuove infezioni da due Vovlas C., Lafortezza R., 1994. Il virus dell'avvizzimento macula- tospovirus (tomato spotted wilt e impatiens necrotic spot) in to del pomodoro su carciofo in Puglia. Informatore Fitopato- Liguria. Informatore Fitopatologico 42: 37-42. logico 44: 42-44. Vaira A.M., Lisa V., Luisoni E., 1992b. Diffusione di due ceppi di Wang M., Gonsalves D., 1990. ELISA detection of various toma- tomato spotted wilt virus in Liguria. Informatore Fitopatologi- to spotted wilt virus isolates using specific antisera to structur- co 42: 59-63. al proteins of the virus. Plant Disease 75: 154-158. Vaira A.M., Lisa V., Dellavalle G., 1992c. Distribution of Weekes R., Barker I., Wood K.R., 1996. An RT-PCR test for the tospovirus in Liguria, Italy. Proc. 5th International Plant Virus detection of tomato spotted wilt virus incorporating immuno- Epidemiology Symposium, Bari 1992, 23-24. capture and colorimetric estimation. Journal of Phytopathology 144: 575-580. Valverde R.A., Can F., Rush M.C., 1993. Yellow mottle of Tomatillo (Physalis ixocarpa) caused by physalis mottle virus. Wijkamp I., Van Lent J., Kormelink R., Goldbach R., Peters D., Plant Pathology 42: 657-660. 1993. Multiplication of tomato spotted wilt virus in its vector, Frankliniella occidentalis. Journal of General Virology 74: 341- Van Os B., Stancatelli G., Mela L., Lisa V., 1993. Ruolo delle pi- 349. ante spontanee ed infestanti nell'epidemiologia di tospovirus in Liguria. Informatore Fitopatologico 43: 40-44. Wilson C.R., 1998. Incidence of weed reservoirs and vectors of tomato spotted wilt tospovirus on southern Tasmanian lettuce Verhoeven J.T.J., Roenhorst J.W., 1994. Tomato spotted wilt farms. Plant Pathology 47: 171-176. virus: ecological aspects in ornamental crops in the Nether- lands from 1989 to 1991. Acta Horticulturae 377: 175-182. Zimmer R.C., Haber S., 1992. First report of tomato spotted wilt virus in Manitoba and of Lathyrus sativus as a host. Plant Dis- Verhoeven J.T.J., Roenhorts J.W., 1998. Occurrence of tospovirus ease 76: 753. in the Netherland. In: Peters D. and Golbach (eds.). Recent

Received 12 July 2003 Accepted 29 September 2003